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Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. LXXXV, No. 1
ANNOTATED LIST OF THE
TYPES OF POLYCHAETOUS ANNELIDS IN THE
MUSEUM OF COMPARATIVE ZOOLOGY
By Olga Hartman
University of Southern California
With Three Plates
CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM
November, 1938
PUBLICATIONS OF THE
MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD COLLEGE
There have been published of the Bulletin, Vols. I to LXXXII, and LXXXV, No. 1 ; of the Memoirs, Vol. I to LIV, No. 1, 2 & 3, and Vol. LV.
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Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. LXXXV, No. 1
ANNOTATED LIST OF THE
TYPES OF POLYCHAETOUS ANNELIDS IN THE
MUSEUM OF COMPARATIVE ZOOLOGY
By Olga Hartman
University of Southern California
With Three Plates
CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM
November, 1938
-^* 7ooloqy 'i>
NOV la 193&
X,
.'D x^
No. 1. — Annotated List of the Types of Polychaetous Annelids in the Museum of Comparative Zoology ^
By Olga Hartman
Bibliographic source, museum catalog numbers and type locality are given for ready reference. Synonyms are enclosed in brackets. New synonyms are preceded by an asterisk, uncertain ones by a question mark. Comments are made only on those types where changes of name are made and which have not been treated since first described.
In some specimens, for example the Ehlers' 1868 types, it is not certain that the specimens designated "types" are actually the holo- types. Also, with several of the Chamberlin, 1920 types, para types are included because it is not certain that the holotypes are still extant. A considerable number of types that have been reported as deposited in the Museum have not been found in the collections. They are not included in this paper. Many of the Ehlers' 1887 types, from Florida and vicinity, are dried or badly disintegrated. Little can be added to what is known of them without further collecting in the type localities.
Two genera of the family SABELLIDAE which are represented by species in considerable confusion, and for which important type materials are in the Museum, are taken up separately at the end of this paper. These genera are Eudistylia Bush and Pseudopotamilla Bush.
Thirty families of polychaetous annelids out of a total of fifty -seven families are represented by types in the Museum. These include 160 species, of which 40 are here regarded as synonyms. Eleven others are questionably synonyms also. This leaves a total of 119 types of valid species.
The following list is arranged alphabetically under family names.
APHRODITIDAE
Aphrodita acuminata Ehlers, 1887, pp. 30-42, pi. 5, figs. 1-5; M.C.Z. nos. 737, 1512, 2082, 2200; Florida.
*[Aphrodita leioseta Chamberlin] 1919b, p. 254; M.C.Z. no. 14; California, is identical with A. japonica Marenzeller. Monro (1933, p. 12) has already commented on the identity of A. leioseta and A.
• I beg to thank the Museum of Comparative Zoology for enabling me to spend two months in Cambridge, in the fall of 1937. Without the opportunity to examine the type material it would have been impossible to settle the various questions of synonymy dealt with in this paper.
4 bulletin: museum of comparative zoology
solitaria Essenberg (1917, p. 408). Both appear to be the same as A. japonica.
Aphrodita ohtccfa Ehlers, 1887, pp. 42-44, pi. G, figs. 1-8; M.C.Z. no. 723; Florida.
Laetmomce nuchipapUlata Augener, 1906, pp. 101-113; pi. 1, figs. 2-7; M.C.Z. no. 2205; West Indies.
Pontogenia maggiae Augener, 1906, pp. 103-106, pi. 1, figs. 8-15; M.C.Z. no. 2127; West Indies.
Pontogenia sericoma Ehlers, 1887, pp. 46-48, pi. 7, figs. 1-5; M.C.Z. no. 818; Havana.
POLYNOIDAE
*[Arctonoe lia Chamberlin] 1920, pp. 6-7, pi. 1, figs. 1-4, pi. 2, figs. 1-3; M.C.Z. paratypes 2190, 2191; Alaska, is identical with A. vittata (Grube). The paratypes compare favorably with specimens of A. vittata collected from the l)ranchial grooves of Cryptochiton and other hosts from California. The name Arctonoe Chamberlin is re- tained because it has priority over Ilalosydnoides Seidler (see also Hartman, on the types of Polynoidae in U.S.N.M., in press, 1937).
'^[Halosydna leioseta Chamberlin 1919a, pp. 2-3; M.C.Z. no. 2140; California, is identical with Arctonoe pulchra (Johnson), new comb. Chamberlin's description, as also the type specimen, agrees in detail with the characters of A. pulchra. This species, like Arctonoe vittata, is a west coast commensal, commonly found with a holothurian, more rarely with other echinoderms.
Halosydna latior Chamberlin, 1919a, pp. 1-2; M.C.Z. no. 2138; California, is readily distinguished by its broadly imbricated, reniform scales, the heavy marginal fringe, and the greatly elongated neph- ridial papillae.
Harmothoii iphionelloides Johnson, 1901, pp. 391-392, pi. 1, figs. 2-7; M.C.Z. no. 1897; Washington.
^[Hesperonoe senilis Chamberlin] 1919b, pp. 252-254, pi. 1, figs. 1-4; M.C.Z. no. 179a; California, is identical with Hesperonoe complanata (Johnson). The scales are pale, ornamented with uniformly dis- tributed, fine prickles. 1 have found the species commensal with Callianassa californiensis Dana, in San Francisco Bay. The closely related //. adventor (Skogsberg) has smooth scales, the body is notably more depressed, and it is commensal with Urechis caupo Fischer and McGinitie.
hartman: polychaetous annelids 0
Lepidametria gigas (Johnson) 1897, pp. 172-175, pi. 7, figs. 33, 42, pi. 8, figs. 48, 49; M.C.Z. no. 964; California.
Lepidonotus citrifrons Augener, 1906, pp. 113-115, pi. 2, figs. 31-38, pi. 3, figs. 39, 40; M.C.Z. no. 2121, 2227, 2228; West Indies.
Lepidonotus Jacteus (Ehlers) 1887, p. 52, pi. 9, figs. 1-7; M.C.Z. nos. 786, 1C96, 2009; Florida (see Seidler, 1924, p. 34).
Lepidonotus leius Chamberlin, 1919a, p. 4; M.C.Z. no. 2142; Cali- fornia.
Lepidonotus setosior Chamberlin, 1919a, pp. 3-4; M.C.Z. no. 2141; California.
Lepidonotus {Thormora) tacniatus (Ehlers) 1887, pp. 51-52, pi. 10, figs. 1-8; M.C.Z. no. 1251; Florida (see Seidler, 1924, p. 92).
Nemidia antiUicola Augener, 1906, pp. 126-128, pi. 3, figs. 53-59; M.C.Z. nos. 2105, 2120; West Indies.
PolynoeUa pachjlepis Augener, 1906, pp. 120-122, pi. 3, figs. 48-52; M.C.Z. nos. 2018, 2232; West Indies.
POLYODONTIDAE
[Euarche tubifex Ehlers] 1887, pp. 54-56, pi. 12, figs. 1-7, pi. 13, fig. 1; M.C.Z. no. 659; Florida, is identical with Eupanthalis kinbergi Mcintosh (see Monro, 1928, p. 568).
SIGALIONIDAE
Psavimolyce flnccifera Augener, 1906, pp. 109-113, pi. 2, figs. 24-30; M.C.Z. nos. 2107, 2113; West Indies.
Sigalion pourtalesii Ehlers, 1887, pp. 57-60, pi. 15, figs. 1-4; pi. 16, 1-10; M.C.Z. nos. 37, 813; Florida. Chamberlin (1919c, p. 91) reported this species from the Gulf of Mexico. Treadwell's record (1914, p. 183) from southern California, is presumably not this species. One speci- men, from Ballast Point, San Diego, so labelled in the collections of the University of California at Berkeley, differs from S. pourtalesii Ehlers in at least the following respects: (1) The superior part of the neuro- podial lobe is drawn out in a point, and (2) the parapodia are propor- tionately longer than in S. pourtalesii.
Sthenelais simplex Ehlers, 1887, pp. 69-63, pi. 13, figs. 2, 3, pi. 14, figs. 1-6; M.C.Z. no. 772; Florida.
Sthenelais verruculosa Johnson, 1897, p. 187, pi. 9, fig. 62, pi. 10, figs. 65-65d; M.C.Z. no. 1939; California north to British Columbia (see Berkeley 1923, p. 216).
6 bulletin: museum of comparative zoology
AMPHINOMIDAE
[Chloeia eughcMs Ehlers] 1887, pp. 18-24, pi. 1, figs. 1, 2, pi. 2, figs. 1-8, pi. 3, figs. 1-4; M.C.Z. nos. 617, 664, 720; Florida, is identical with C. viridis Schmarda (see Monro, 1933, p. 9).
EUPHROSINIDAE
Euphrosine triloba Ehlers, 1887, pp. 31-33, pi. 4; M.C.Z. no. 1250; Florida.
PHYLLODOCIDAE
[Phyllodoce ocidata (Ehlers)] 1887, pp. 135-140, pi. 40, figs. 4-6; M.C.Z. no. 795; Florida, is an Anaiiides (see Bergstrom, 1914, p. 144), identical with Lepadorhynchus erythrophyllus Schmarda (see Augener 1925, pp. 23-24). Schmarda's type came from Jamaica.
Paranaitis papiUosa (Ehlers) 1887, pp. 140-142, pi. 40, figs. 7-9; M.C.Z. no. 839; Florida, was originally described as Anaitis.
Steggoa gracilior Chamberlin, 1919a, pp. 5-6; M.C.Z. no. 2144; California.
HESIONIDAE
Castalia hcsionidcs Augener, 1906, pp. 155-157, pi. 6, figs. 106-109; M.C.Z. no. 2012; West Indies.
[mesioneviUigera Ehlers], 1887, pp. 143-147, pi. 41, figs. —6; M.C.Z. no. 837; Florida, as also H. praetcxta Ehlers, (see below) is seemingly identical with //. protochuna Schmarda. Ehlers has already suggested this possibility in his original description.
[Hesione praetexia Ehlers] 1887, p. 147; M.C.Z. no. 762; Florida, is identical with H. vittigera. They were separated on the basis of color. Ehlers has already discussed their possible identity.
SYLLIDAE
Branchiosyllis oculata Ehlers, 1887, pp. 148-151, pi. 39, figs. 1-7; M.C.Z. no. 844; Florida.
CampesylUs minor Chamberlin, 1919a, p. 10; M.C.Z. no. 2151; California.
Hcsperalia californiensis Chamberlin, 1919a, p. 9; M.C.Z. no. 2149; California.
hartman: polychaetous annelids /
Hesperalia nans Chamberlin, 1919a, pp. 9-10; M.C.Z. no. 2150; California. This may represent the epitokous stage of Hesperalia cali- fornierisis Chamberhn. The latter is one of the commonest of Syllidae in the littoral zone of southern California.
Pionosyllis elongata Johnson, 1901, pp. 403-405, pi. 6, figs. 67—70, pi. 7, fig. 71; M.C.Z. no. 1937 (cotype); Washington. Common in the littoral zone of the northeast Pacific. Berkeley (1938, p. 41) has en- hanced the description and rendered it distinguishable from that of nearly related species.
Pionosyllis lucida Chamberlin, 1919a, p. 8; M.C.Z. no. 2180 California.
Pionosyllis pigmentata Chamberlin, 1919a, p. 7; M.C.Z. no. 2148 California.
Trypanosyllis vittigera Ehlers, 1887, pp. 151-154, pi. 40, figs. 1-3 M.C.Z. no. 843; Florida.
Typosyllis hella Chamberlin, 1919a, p. 7; M.C.Z. no. 2147; Cali- fornia.
NEREIDAE
*?Ceratonereis articulata (Ehlers) 1887, pp. 114-116, pi. 36, figs. 1-4; M.C.Z. no. 764; Florida, was first described as a Nereis. It is char- acterized by an absence of paragnaths on the maxillary ring. The teeth of the oral ring were described as pale, obtuse-conical, forming a simple series. The type is tiny, incomplete, proboscis is lacking, and it has been dried at some time. It may be a young stage of another described species.
*[Nereis bicruciata Augener] 1906, pp. 151-153, pi. 5, figs. 102-104, pi. 6, fig. 105; M.C.Z. no. 2277; West Indies, is identical with Nereis nigripes Ehlers, which is, in turn, the same as N. riisei Grube. Augen- er's type is considerably the smaller of the two. The feet have the typical black patches, distributed as in A^. nigripes and the parapodial lobes of the two are strikingly similar. The difference in the dentition of the types of Augener and Ehlers, as originally described, is : Area VI of the proboscis has 6 or 7 teeth in a heap in A^. nigripes and only 4 or 5 in iV. bicruciata, areas III and IV have fewer teeth in A^. bicruciata. Both of these differences are not unlikely in specimens of different ages or sizes. More recently Treadwell (1931, p. 14) has described Nereis decora, from Brazil. The description agrees with that for N. riisei Grube.
8 bulletin: museum of comparative zoology
Nereis latesccns Chamberlin, 1919a, pp. 10-11; M.C.Z. no. 152: California.
Nereis mediator Chamberlin, 1919a, p. 11; M.C.Z. no. 2153; Cali- fornia.
Nereis mendocinana Chamberlin, 1919b, pp. 256-257, pi. 1, fig. 5; M.C.Z. no. 2129; California, is separable from other species of Nereis in its dentition. Areas 1, 111 and V of the proboscis lack teeth, area VI has 7 small cones in a heap on the left side and 6 on the right side. Areas Vll and VIII have a single continuous series of 7 larger teeth on the maxillary side and a series of progressively smaller teeth toward the oral side. Jaws have 7 (to 9) quadrate teeth on the cutting edge.
[Nereis nigripes Ehlers] 1868, pp. 508-510; M.C.Z. no. 54; Florida, Haiti; is Nereis riisei Grube (see Monro, 1933, pp. 43-44, for further synonymy and discussion).
'^•[Nereis versipcdata Ehlers] 1887, pp. 116-117, pi. 36, figs. 5-10; M.C.Z. no. 764; Florida, is a Ceratonereis, identical with C. irritabilis (Webster). I have examined Webster's type in the U.S. National Museum and compared it with the original illustrations. These give an accurate portrayal of the diagnostic parts. The type of N. versipcdata has numerous paragnaths on area III (unpaired) contrary to the con- dition illustrated. It is thus as found in C. irritabilis.
Nereis procera Ehlers, 1868, pp. 557-559, pi. 23, fig. 2; M.C.Z. no. 155; Gulf of Georgia, has numerous tiny paragnaths on both oral and maxillary rings of the proboscis. In the posteriormost parapodia the dorsal lobe is flattened, equitriangular, the dorsal cirrus attached near its proximal base and extends distally beyond the dorsal lobe. Middle and ventral parapodial lobes are small, digitiform, subequal; the neuroacicular lobe is truncate; ventral cirrus tiny.
Perinercis floridana (Ehlers) 1868, pp. 503-506; M.C.Z. no. 119; Florida, has been redescribed by Langerhans (1879, pp. 280-290) and others.
Uncinereis agassizi (Ehlers) 1868, pp. 542-546, pi. 23, fig. 1; M.C.Z. no. 179; California, includes Nereis notomacula Treadwell (1914, pp. 191-192, pi. 11, figs. 8-12; California), Uncinereis subita Chamber- lin (1919c, pp. 215-219, pi. 30, figs. 1^; California) and Platijncreis kohiensis Mcintosh (1885, pp. 210-212, pi. 34, figs. 3-6, pi. 16a, figs. 2-4; Japan).
NEPHTYIDAE
(Nephtys Cuvier is used instead of the more widely used Nephthijs Savigny because the former has priority. See Hartman, 1938b, p. 145).
hartman: polychaetous annelids 9
Nephtys bucera Ehlers, 1868, pp. 617-619, pi. 23, fig. 8; M.C.Z. no. 209; Massachusetts Bay, consists of over 100 segments. The type is in two pieces, with some segments perhaps missing. It is much like N. incisa Malmgren from Europe. The proboscis has 22 rows of papillae, acicular lobes are conical, the recurved cirri are cirriform, the dorsal cirrus smaller than the recurved cirrus. In the posterior part of the body, however, the postsetal lobes of both rami are expanded, folia- ceous (pi. 1, fig. 1) instead of reduced as in N. incisa.
Nephtys cirrosa Ehlers, 1868, pp. 624-626, pi. 23, figs. 6, 37, 38; M.C.Z. no. 1242; England.
Nephtys discors Ehlers, 1868, pp. 626-629, pi. 23, figs. 39-10; M.C.Z. nos. 127, 700; Maine, has dorsal and ventral postsetal lobes large, foliaceous throughout, but largest in the anterior region (pi. 1, fig. 3.). Dorsal cirri are short, never as large as the recurved cirri except in the first and last few segments; recurved cirri are first present from the sixth segment as minute papillae; they are fleshy digitiform, slightly recurved where best developed, but never extending much beyond the dorsal half of the foot. Notoacicular lobes are bifid through the anterior half, then gradually become entire (pi. 1, fig. 2).
Nephtys inermis Ehlers, i887, pp. 125-128, pi. 38, figs. 1-6; M.C.Z. no. 1088; Florida.
Nephtys phyllocirra Ehlers, 1887, pp. 131-134, pi. 38, figs. 7-11; M.C.Z. nos. 62, 1033; Florida.
*[Nephtys pida Ehlers] 1868, pp. 632-635, pi. 23, figs. 9, 35; M.C.Z. nos. 246, 830; New England, is close to, if not identical with, N. caeca (Fabricius). Recurved cirri are first present at the fourth setigerous segment; acicular and setal lobes agree with those typical of N. caeca and the proboscis is proximally covered with coarse, low papillations.
Nephtys squamosa Ehlers, 1887, pp. 128-131, pi. 37, figs. 7-10; M.C.Z. nos. 735, 740, 741, 836; Florida.
GLYCERIDAE
*[GJycera exigua Chamberlin] 1919a, pp. 13-14; M.C.Z. no. 2158; California, is identical with Glycera viacrohranchia Moore (1911, pp. 301-302; California). Chamberlin's type is smaller than is typical of adults of this species, but it agrees in details of structure.
Glycera nana Johnson, 1901, p. 411, pi. 10, figs. 103-1 03a; M.C.Z. no. 1878; Puget Sound, Washington.
tGlycera oxycephala Ehlers, 1887, pp. 121-123, pi. 41, figs. 7-11;
10 bulletin: museum of comparative zoology
M.C.Z. no. 713; probably West Indies, may be identical with Glycera gigajitca Quatrefages, from Europe.
Glycera rohnsta Ehlers, 1868, pp. 656-658, pi. 24, figs. 31, 32; M.C.Z. no. 193; California.
Glycera rugosa Johnson, 1901, pp. 409-411, pi. 10, figs. 101-102; M.C.Z. no. 2296; Puget Sound, is nearly related to Gl. americana Leidy, from the east coast of North America. The two may differ slightly in the shape of the setal lobes. Glycera corrugata Baird (1863, p. 109; Vancouver) may be identical. Its type, if still extant, should be reinvestigated. Glycera peruviana Quatrefages (1865, p. 177) from Peru, is another nearly related form.
GONIADIDAE
Goniada eremita var. quinquelabiata Augener, 1906, p. 158; M.C.Z. no. 2297; West Indies, is represented in the collections by a single specimen. Though not designated type, the locality is as stated in the original description.
EUNICIDAE
Eunice collini Augener, 1906, pp. 133-135, pi. 4, figs. 66-73; M.C.Z. no. 2011; West Indies.
Eunice floridana (Pourtales) 1869, p. 108; M.C.Z. nos. 665, 666, 671-675; Florida, has been redescribed and illustrated by Ehlers, 1887, pp. 88-90, pi. 22, figs. 1-7.
[Ewiice fucata Ehlers] 1887, pp. 91-93, pi. 25, figs. 8-20; M.C.Z. nos. 124, 151, 198, 242, 724; Florida, is identical with E. schemacephala Schmarda (see Augener, 1925, p. 28).
[Eu7iice antillensis Ehlers] 1887, pp. 84-85, pi. 24, figs. 5-7; M.C.Z. nos. 140, 799, 810; Florida, is identical with Eunice longicirrata Webster (Monro, 1930, p. 121; 1933, pp. 61-62).
[Eunice conglomerans Ehlers] 1887, pp. 93-95, pi. 23, figs. 1-9, pi. 24, figs. 1-4; M.C.Z. no. 826; Florida, is identical with Eunice filamentosa Grube (see Augener, 1925, p. 9).
Eunice tibiana (Pourtales) 1869, pp. 108-109; M.C.Z. nos. 719, 743, 745; West Indies; is redescribed by Ehlers, 1887, pp. 90-91, pi. 27, figs. 1-13.
[Eunice violaceo-maculata Ehlers] 1887, pp. 86-87, pi. 24, figs. 11, J2, pi. 25, figs. 1-7; M.C.Z. nos. 130, 163, 657; Florida, is considered identical with Eunice rousseaui Quatrefages (see Fauvel, 1923, p. 403).
hartman: polychaetous annelids 11
*[Eunice valens Chamberlin] 1919b, pp. 257-258, pi. 1, figs. 6-8; M.C.Z. nos. 120, 121; California. This was incorrectly included under E. biannulata Moore (see Hartman 1938a, p. 96). The latter was therein designated E. longicirrata var. Eunice valens and E. biannulata differ in parapodial and setal structures.
[Lysidice no^a Ehlers] 1887, pp. 100-102, pi. 30, figs. 1-9; M.C.Z. nos. 711, 794; Florida, was placed in synonymy with L. brachycera Schmarda by Augener (1925, p. 29) who considers both the same as L. ninetta Audouin and Edwards.
Nicidion brevis Ehlers, 1887, pp. 98-99, pi. 28, figs. 9-14; pi. 29, figs. 1, 2; M.C.Z. no. 833; Florida.
Paramarphysa lo7igu1a Ehlers, 1887, pp. 99-100, pi. 29, figs. 3-12; M.C.Z. no. 879; Havana.
Rhamphobrachium agassizi Ehlers, 1887, pp. 70-73, pi. 17, figs. 1-5, pi. 18, figs. 1-9; M.C.Z. nos. 754, 779, 788, 789; Florida.
ONUPHIDAE
Diopatra pourtalesii Ehlers, 1879, p. 273; M.C.Z. nos. 687, 785, 815, 874; Florida, was redescribed by Fillers, 1887, pp. 74-75, pi. 19, figs. 6-10; pi. 20, figs. 1-6.
Nothria elegans Johnson, 1901, pp. 406-407, pi. 8, figs. 77-85; M.C.Z. no. 1895; Washington.
Nothria iridescens Johnson, 1901, p. 408, pi. 8, figs. 86, 87, pi. 9, figs. 88-92; M.C.Z. no. 1887; British Columbia.
Paradiopatra fragosa Ehlers, 1887, pp. 75-76, pi. 20, figs. 7-14, pi. 21, figs. 1-4; M.C.Z. nos. 656, 687, 785, 815, 874; Florida.
Paradiopatra glutinatrix Ehlers, 1887, pp. 76-78, pi. 18, figs. 10-15; pi. 19, figs. 1-5; M.C.Z. nos. 2337-2351; Florida.
Paronuphis gracilis Ehlers, 1887, pp. 78-82, pi. 21, figs. 5-13; Florida.
LUMBRINERIDAE
t^Aracoda debilis Ehlers, 1887, p. 113, pi. 35, figs. 5-8; M.C.Z. no. 757; Florida, is a Drilonereis. Parapodia are provided with stout acicular setae and jaw pieces have attenuated carriers. The forceps (maxillae I) have a few teeth on their inner, basal margin. This species may be identical with Drilonereis filum Claparede, reported from both sides of the Atlantic and Pacific oceans.
12 bulletin: museum of comparative zoology
[Arabella rnimctica Chamberlin] 1919a, pp. 12-13; M.C.Z. no. 2156; California, is identical with Arabella iricolor (Montagu) (see Berkeley, 1932, p. 313).
[Arabella muHidentaia Ehlers] 1887, pp. 112-113, pi. 34, figs. 8, 9, pi. 35, figs. 1-4; M.C.Z. no. 825; Plorida, is identical with Arabella iricolor (Montagu) (see Fauvel, 1923, p. 438).
""[Arabella munda Chamberlin] 1919b, pp. 258-259; M.C.Z. no. 211; California, is identical with Arabella semimaculata Moore, 1911, pp. 295-297, pi. 20, figs. 143-149. A characteristic feature is the fleshy, digitate, obliquely directed parapodial lobe.
Biborin ecbola Chamberlin, 1919a, p. 13; M.C.Z. 2157; California.
Lumbrineris bidens Ehlers, 1887, pp. 103-104, pi. 31, figs. 7-17; M.C.Z. nos. 750, 756, 760; Florida.
*[Lumbrineris floridana Ehlers] 1887, p. 103, pi. 30, figs. 10-15; M.C.Z. nos. 838, 841 ; Florida, is identical with Lumbrineris latreilli Audouin and M. Edwards. The prostomium is broadly rounded in front (sugar-loaf shaped), not conical as shown by Fauvel (1923, fig. 171 m).
Lumbrineris robusta Ehlers, 1887, pp. 104-105, pi. 31, figs. 1-6; M.C.Z. nos. 742, 747; Florida and Havana.
"[Lumbrineris zonata Johnson] 1901, pp. 408-409, pi. 9, figs. 93-100; M.C.Z. no 1954; Puget Sound, is probably identical with Lumbrineris sarsi Kinberg. The identity of this common eastern Pacific species with L. impatiens Claparede seems likely. Kinberg's type is from Ecuador. I have compared specimens from Alaska south to Lower California and believe them all the same. Specimens are commonly encountered in sand under rocks in the intertidal zone, and in root masses of eel grasses.
Ninoe kinbergi Ehlers, 1887, pp. 105-106, pi. 32, figs. 1-9; M.C.Z. no. 881 ; Florida.
STAURONEREIDAE
Stauronereis gregarious Mayer, 1900, pp. 1-14, pi. 1-3; M.C.Z. nos. 1797, 1798; Florida.
[Stauronereis gregarious Mayer], 1900, pp. 1-14, pi. 1-3; M.C.Z. nos. 1797, 1798; Florida, was later identified with Eunioe fuoata Ehlers, by Mayer, 1902. The latter, however, has been shown to be Eunioe soheviacephala Schmarda (see Augener, 1925, p. 29).
hartman: polychaetous annelids 13
ORBINIIDAE
Nainereis hespera Chamberlin, 1919a, pp. 14-15; M.C.Z. no. 2161; California, differs from Nainereis laevigata (Grube) in having black parapodial patches and branchiae from the thirteenth or fourteenth segments.
Nainereis nannohranckia Chamberlin, 1919b, pp. 260-261, pi. 2, fig. 10, pi. 3, fig. 1; M.C.Z. no. 2136; California, differs from Naiyiereis laevigata (Grube) in having no branchiae on the anterior thirty seg- ments. Also, the branchiae are short where best developed, hardly meeting in the mid-dorsal line. Both this and N. hespera may prove to be only variations and color varieties of N. laevigata Grube, which is a common species in the northeast Pacific.
*Scoloplos acmeceps Chamberlin, 1919a, pp. 15-16; M.C.Z. no. 2162; California, is identical with Scoloplos elongata Johnson (1901, pp. 412- 413, pi. 10, figs. 105-110; Puget Sound). The latter name is preoccu- pied by Quatrefages, 1865, hence Chamberlin's name replaces it. A typical parapodium (30th) is shown in pi. 1, fig. 4.
SPIONIDAE
Anaspio boreus Chamberlin, 1920, p. 18, pi. 4, figs. 2-4; M.C.Z. no. 2523; Alaska.
Scolecolepides arcticus Chamberlin, 1920, pp. 17-18, pi. 3, figs. 5-7, pi. 4, fig. 1; M.C.Z. paratypes 2194, 2195; Alaska.
*[Spio minus Chamberlin] 1920, p. 16, pi. 3, figs. 1-4; M.C.Z. para- type 2522; Alaska, may be identical with Spio filicornis (Miiller). The latter has been reported from both north Atlantic and north Pacific waters (Berkeley, 1936, p. 475). Chamberlin separated S. mimus on differences observed in posterior parapodia and in the distribution of crotchets. The variations, however, appear to fall within the range found in <S. filicornis.
CIRRATULIDAE
*[Cirratulus cingulatus Johnson] 1901, pp. 422-423, pi. 14, figs. 145- 148; M.C.Z. no. 1906; Washington, agrees with descriptions of C. cirratus (Miiller). The prostomium has paired series of eye spots, setae and parapodial arrangements are as typical for C. cirratus.
Cirratulus cxuberans Chamberlin, 1919b, pp. 263-264; M.C.Z. no. 1285; Panama.
Cirratulus rohustus Johnson, 1901, p. 423, pi. 14, figs. 149, 150; M.C.Z. no. 1882; Puget Sound, Washington.
14 bulletin: museum of comparative zoology
FLABELLIGERIDAE
Wlabelligera haerens Chamberlin, 1919a, p. 16; M.C.Z. no. 2163; California, is represented by a single, small individual (13 mm. long). It may be a young specimen of Flabelligera affinis Sars, as Monro has suggested (1933, p. 4).
Flabelligera infundibular is Johnson, 1901, p. 417, pi. 12, figs. 124- 127; M.C.Z. nos. 2503, 2508; Puget Sound, differs from H. affinis Sars in the distribution of its composite setae and the nature of its sensory papillae.
Stylarioides collarifer Ehlers, 1887, pp. 161-164, pi. 43, figs. 2-7, M.C.Z. nos. 775, 778, 784; Florida.
Stylarioides papillata Johnson, 1901, p. 416, pi. 12, figs. 122, 123; M.C.Z. no. 1900; Washington, is close to, if not identical with, S. plumosa (O. F. Miiller). Monro (1933b, p. 1058), has already pointed out this similarity.
SCALIBREGMIDAE
Kebuita glabra (Ehlers) 1887, pp. 169-172, pi. 45, figs. 1^; M.C.Z. no. 695; Havana, originally described as a Eumenia, was made the type of the genus Kebuita Chamberlin (1919c, p. 390). It agrees in its details with the description of Oncoscolex heterochaetus Augener (1906, pp. 159-161, pi. 6, figs. 110-112) for which Chamberlin erected the genus Gmtsitoa (1919c, p. 390) . The first three segments in the type are slightly set off from the rest of the body by having more conspicu- ous parapodia, provided with sickle-shaped setae (as illustrated by Augener). The dorsal surface of E. glabra is somewhat corrugated as described by Augener. Forked setae were not observed in 0. heter- ochaetus, but Augener suggested the likelihood that they had been broken off or lost. Both have an anal funnel terminating in a crenulate collar.
Augener has already pointed out the possibility that these two names refer to the same species (1906, p. 161). Because of the similar- ity of Ehlers' type with Augener's description, and the proximity of type localities (Havana, 175 fms. and St. Vincent, 424 and 785 fms. respectively) the two are considered identical. Gwasitoa Chamberlin thus becomes a synonym of Kebuita Chamberlin. A revised diagnosis of Kebuita follows.
Body maggot-like; prostomium with lateral processes (frontal horns) ; no eye spots or visible nuchal organs. Parapodia biramous, the
hartman: polychaetous annelids 15
rami well separated from each other, but poorly developed, most con- spicuous m the anterior region, the first three setigerous segments slightly set off from those more posteriorly by their more prominent feet. Setae of first three setigerous segments sickle-shaped; setae in more posterior parapodia capillary and forked. Dorsal and ventral cirri absent. Anal funnel cylindrical, terminating posteriorly in a crenulate collar with short papillae on the ventral edge but without anal cirri.
Remarks. Differs from Asderocheilus Ashworth in lacking anal cirri and in having an anal funnel; differs from Oncoscolex Schmarda (which ma}^ be identical with Hyhoscohx Schmarda, according to Augener, 1923, p. 83) in lacking eye spots, in having parapodia much more reduced, in having an anal funnel without cirri.
The type of the genus is Eumenia glabra Ehlers.
CAPITELLIDAE
Dasi/branchus lumulatus Elilers, 1887, pp. 174-177, pi. 45, figs. 5-9; M.C.Z. no. 847; Florida.
Notomastus angulatus Cliamberlin, 1919a, pp. 16-17; M.C.Z. no. 2164; California.
MALDANIDAE
Clymene cinguJata Ehlers, 1887, pp. 185-188, pi. 47, figs. 2-5; M.C.Z. no. 541; Florida, was made the type of Clymenopsis Verrill (1900, p. 654). It is known through a single anterior end, characterized as follows: Head plaque reduced to a thickened area bearing a pair of parallel nuchal organs; fourth segment with a ventrally produced collarette; stout acicular spines on first three segments; uncini in single rows.
Clymene cirrata Ehlers, 1887, pp. 182-185, pi. 46, figs. 10-13; M.C.Z. no. 2402; Florida, has been dried and is considerably frag- mented. Arwidsson (1907, p. 145) has shown that it is a Leiochone.
Maldane collariceps Augener, 1906, pp. 163-167, pi. 6, figs. 114-120; M.C.Z. 2029; West Indies, is an Asychis. Its affinities with A. biceps (Sars) and A. coronaia (Moore) have been discussed by Augener.
Maldane cucidligera Ehlers, 1887, pp. 178-182, pi. 46, figs. 1-9; M.C.Z. no. 771; Florida.
Nicomache personata Johnson, 1901, pp. 419-420,' pi. 13, figs. 134- 139; M.C.Z. no. 1918; Washington.
16 bulletin: museum of comparative zoology
Nichomachella (?) pida Ehlers, 1S87, pp. 192-193, pi. 47, fig. 1; M.C.Z. no. 241; Florida, is a Praxillura Verrill (Arwidsson, 1907, p. 26).
OWENIIDAE
[Ammochares occidentale Johnson] 1901, pp. 420-421, pi. 14, figs. 140-142; M.C.Z. no. 1997; Washington; is identical with Owenia fusiforviis (delle Chiaje) (see Berkeley, 1930, p. 67).
SABELLARIIDAE
[Idanthyrsus ornamentatus Chamberlin] 1919b, pp. 262-263, pi. 3, figs. 2-5; M.C.Z. no. 156; California, is identical with Idanthyrsus johnstoni (Mcintosh) (see Berkeley, 1930, p. 74).
Lygdamis asteriformis (Augener) 1906, pp. 167-171, pi. 7, figs. 129-138; M.C.Z. nos. 2053, 2054; West Indies (see Johansson, 1927, p. 86).
Sabellaria nanella Chamberlin, 1919b, pp. 261-262, pi. 2, figs. 5-7; M.C.Z. paratype no. 482; California, difi'ers from other described species in its opercular spines. Those of the outer, middle and inner sets are illustrated in pi. 3, figs. 10, 8 and 9 respectively.
PECTINARIIDAE
Pcctinaria (Petta) peUucida Ehlers, 1887, pp. 194-199, pi. 44, figs. 1-9; M.C.Z. no. 2559; Florida, is represented by a unique specimen not designated type, but presumably labelled in Ehlers' handwriting, and collected from 270 fms. in the Santaren Channel.
AMPHARETIDAE
Ampharete rcducta Chamberlin, 1920, p. 24, pi. 5, figs. 4, 5; M.C.Z. paratype no. 2192; Alaska.
Avipkidcis nasuta Ehlers, 1887, pp. 232-236, pi. 49, figs. 1-6; M.C.Z. no. 755; Florida, has been assigned to Anobothnis (see also Hessle, 1917, p. 107).
Ampkidcis proccra Ehlers, 1887, pp. 226-232, pi. 48, figs. 1-9; M.C.Z. no. 852; Florida.
Auchenoplax crinita Ehlers, 1887, pp. 209-214, pi. 44, figs. 10-16; M.C.Z. nos. 850, 873; Florida.
hartman: polychaetous annelids 17
Melinna monocera Augener, 1906, pp. 177-180, pi. 6, figs. 121-125 M.C.Z. no. 2128; West Indies.
Melinna parumdentata Ehlers, 1887, pp. 214-220, pi. 50, figs. 1-9 M.C.Z. no. 872; Florida.
Sabellidcs anops Johnson, 1901, pp. 424^25, pi. 15, figs. 157-161 pi. 16, figs. 162, 163; M.C.Z. no. 1892; Washington (?) is an Amage Berkeley (1929, p. 311) considers it likely that this is the same as A auricula Malmgren, known from the boreal Atlantic and Pacific.
Schistocomus hiltoni Chamberlin, 1919a, pp. 17-18; M.C.Z. no. 2166; California.
TEREBELLIDAE
Amphitrite rob usta Johnson, 1901, pp. 425-426, pi. 16, figs. 164-168; M.C.Z. nos. 954, 2392; Puget Sound, has been assigned to Neoamphi- iriie Hessle (1917, p. 184).
Amphitrite spiralis Johnson, 1901, pp. 426^27, pi. 16, figs. 169- 171; M.C.Z. no. 1874; Washington, is a Neoleprea Hessle (1917, p. 193).
Terchella bruneo-comata Ehlers, 1887, pp. 237-241, pi. 51, figs. 1-5; M.C.Z. no. 831; Florida, was made the type of the subgenus Am- phitrides Augener (1922, p. 47). Monro (1933, p. 1071) has reported it from the Galapagos Islands and consideres Terebella panamena Chamberlin, from Panama, identical.
Eupolymnia crescentis Chamberlin, 1919b, pp. 265-266, pi. 3, figs. 6, 7; M.C.Z. no. 2135; California, is distinct from E. nebulosa (Mon- tagu). The ventral thoracic plates, save those of the first three seg- ments which are broadly rectangular, are broad, trapezoidal, with the anterior, longer side slightly arcuate. The scutes are broadest in the regions of the seventh to ninth segments, here they are about five times as broad as long. They become rapidly smaller from the twelfth seti- gerous segment, the fifteenth is tiny, the last two thoracic segments lack scutes. The cephalic lobe has numerous minute eye spots, but the lobe is not conspicuously elongate as in E. nebulosa. Thoracic un- cinigerous tori are proportionately longer and extend into the angle formed by the lateral extensions of the thoracic plates. Thoracic uncini have a more elongate base, with tooth nearer the base of the hook (see Chamberlin, 1919b, pi. 3, figs. 6, 7).
Leaena videns Chamberlin, 1919a, p. 18; M.C.Z. no. 2167; Cali- fornia.
18 . bulletin: museum of comparative zoology
Naneva hespera Chamberlin, 1919a, p. 19; M.C.Z. no. 2169; Cali- fornia.
Pista hrevihranchia Chamberlin, 1919b, pp. 264-265; pi. 2, figs. IHI; M.C.Z. no. 502; California, is characterized by having the lateral thoracic lappets continued through 5-7 anterior segments (see also Moore, 1923, p. 196). Chamberlin's name is pre-occupied by Caullery (1915, p. 76). The name P. brembranchiata, apparently inadvertently used by Moore (1923, p. 196) is suggested to replace P. brevibranchia Chamberlin.
Pista fratrella Chamberlin, 1919a, pp. 18-19; M.C.Z. no. 2168; California.
Scionides dux Chamberlin, 1919b, pp. 206-207, pi. 3, fig. 9; M.C.Z. no. 2134; California.
Terebella cetrata Ehlers, 1887, pp. 248-253, pi. 52, figs. 9-14; M.C.Z. no. 834; Florida, is a Pista (see Hessle, 1917, p. 163).
Tcrehclla reticulata Ehlers, 1887, pp. 245-248, pi. 52, figs. 5-8; M.C.Z. no. 840; Florida, has been dried and is badly broken. It may be a Neoamphitrite (see also Hessle, 1917, p. 184). Thoracic noto- podial setae are distally denticulate.
[Terebella turgidula Ehlers] 1887, pp. 241-245, pi. 52, figs. 1^; M.C.Z. no. 846; Florida, is a Eupolymnia (see Hessle, 1917, p. 178), identical with E. crassicornis (Schmarda) (see Augener, 1925, p. 36).
SABELLIDAE
Bispira poly morpha Johnson, 1901, pp. 428-430, pi. 17, figs. 179-183, pi. 18, figs. 184, 185; M.C.Z. nos. 1901, 1902, 1903, 1941; Puget Sound to Pacific Grove, includes four species, — Eudistylia polymorpha, E. vancouveri (Kinberg) and Schizobranchia spp. (see p. 26 for discussion).
Branchiomma bioculatum Ehlers, 1887, pp. 260-263, pi. 53, figs. 1-9; M.C.Z. no. 669; Florida, is a Megalomma (Johansson, 1927, p. 132). It should be examined in the light of other collections from the type locality.
Branchiomma lobifcriim Ehlers, 1887, pp. 254-259, pi. 53, figs. 10-15; M.C.Z. no. 835; Florida, is a Megalomma (see Johansson, 1927, p. 132). It has a straight collar, the dorsal lobes are inconspicuous, most of the ventral radioles have terminal eyes, and the branchiae are greatly elongated.
Chone ungavana Chamberlin, 1920, pp. 26-27, pi. 6, figs. 1-4; M.C.Z. paratype no. 2550; Alaska.
[Dasychone conspersa Ehlers] 1887, pp. 266-270, pi. 54, figs. 1-6;
hartman: polychaetous annelids 19
M.C.Z. no. 848; Florida, is identical with Branchiomvia nigromaculata (Baird) (see Johansson, 1927, pp. 162-163).
[Distylia monterea ChamberHn] 1919b, pp. 267-268; M.C.Z. no. 1941, is identical with Eudistylia polymorpha, see p. 26 for discussion.
Hypsicomus circumspiciens Ehlers, 1887, pp. 271-277, pi. 55; figs. 5-13, pi. 56, figs. 1-3; M.C.Z. no. 662; Florida, was made the type of the genus Hypsicomatopsis Augener. It is likely that Protulides elegans Webster (1884, pp. 325-326, pi. 11, figs. 63-74; Bermuda) is the same as H. circumspiciens.
Megachone aurantiaca Johnson, 1901, pp. 430-431, pi. 18, figs. 186- 192; M.C.Z. no. 1933; Washington.
*[Myxicola monads ChamberHn] 1919a, p. 20; California, is identical with Myxicola infundibulum (Renier), see discussion under M. pacifica.
*[Myxicola pacifica Johnson] 1901, pp. 431-432, pi. 19, figs. 193-198; M.C.Z. nos. 1879, 1893; Washington, is identical with M. infundihuluvi (Renier). This species is known from both north Atlantic and Pacific oceans. Individuals from British Columbia south to southern Cali- fornia, agree in important details with descriptions given for Atlantic species. Characteristic features of a specimen from California, are shown in pi. 1, figs. 5-11, pi. 2, fig. 1.
Potamcthus spathiferus (Ehlers) 1887, pp. 278-283, pi. 54, figs. 7-11, pi. 55, figs. 1^; M.C.Z. no. 800; Florida.
*[Potamilla clara ChamberHn] 1919a, p. 20; M.C.Z. no. 2171; Cali- fornia, is a Megalomma, and has affinities with M. roulei (Gravier). M. clara has one pair of dark, compound terminal eyes; the thoracic collar lacks dorsal lappets.
^[PotamiUa colorata ChamberHn] 1919a, p. 21; M.C.Z. no. 2173; California, is identical with Megalomma clara (see above). Two of the radioles have dark, compound eyes. Collar lacks dorsal lappets.
'^[Pseudopotamilla lampra ChamberHn] 1919a, p. 22; M.C.Z. no. 2176; California, is identical with Pseudopotamilla occelata Moore, see p. 506 for discussion.
*[Pseudopotamilla macrops ChamberHn] 1919a, p. 22; M.C.Z. no. 2177; California, is identical with Pseudopotamilla intermedia Moore, see p. 506 for discussion.
^[Pseudopotamilla panamaica ChamberHn] 1919b, pp. 268-269, pi. 3, fig. 8; M.C.Z. no. 72; Panama, is a Megalomma, identical with M. vesiculosum (Montagu) cosmopolitan. Terminal eyes are numerous; dorsal collar lappets are long, foliaceous; lateral collar lobes are con- spicuous.
*[Pseudopotamilla parva ChamberHn] 1919a, p. 21; M.C.Z. no. 2175;
20 bulletin: museum of comparative zoology
California, is doubtfully referred to Mcgalonuna. The type resembles P. clara, except that terminal eyes cannot be made out. Only a few anterior segments are present. Dorsal collar lobes are greatly reduced.
^[Pscudojjotamilla yauroys Chamberlin] 1919a, p. 21; M.C.Z. no. 2174; California, is identical with P. occdata Moore. See p. 507 for dis- cussion.
^[PseudojMtamilla scotia Chamberlin] 1919a, p. 22; M.C.Z. no. 2178; California, is identical with P. occdata Moore. See p. 507 for discussion.
SERPULIDAE
Crucigera hespera Chamberlin, 1919b, p. 270, pi. 2, fig. 9; M.C.Z. no. 164; California.
Crucigera zygoj:)hora (Johnson) 1901, pp. 433-434, pi. 19, figs. 205- 208; M.C.Z. no. 1875; Washington, has been subsequently treated by Bush (1904, p. 233).
'lEupomatus inicrcans Chamberlin, 1919a, p. 23; M.C.Z. no. 2178; California, is probably identical with Eupomatus uncinatus Philippi, as already suggested by Chamberlin. The unique type has been dried.
F'Uograna kuxhyi Ehlers, 1887, pp. 314-320, pi. 56, figs. 4-8; M.C.Z. no. 739; Florida.
Ilyalopomatus langerhansi Ehlers, 1887, pp. 304-307, pi. 60, figs. 10-15 ; M.C.Z. nos. 688, 759 ; Havana.
Phicostegus incomptus Ehlers, 1887, pp. 300-304, pi. 60, figs. 1, 2; M.C.Z. no. 811; Havana.
Protuhi anicmmia Ehlers, 1887, pp. 321-324, pi. 59, figs. 4-9; M.C.Z. no. 808 ; Florida. The type jar contains several pieces of one specimen and parts of two tubes, the latter presumably belonging to the species. Augener (1925, p. 39) concluded that this may be identical with P. longisda Schmarda (1861, p. 32) from Jamaica. This synonymy seems doubtful, however, for several reasons. The tube of P. antcnnata is externally ridged and not lined with a chitinous layer; the uncini and collar are different. I am unable to add to Ehlers' description of the collar. The unique type has been dried and is broken. A few of the posterior setae are longer than indicated by Ehlers, but none is con- spicuously long.
Protula suhmedia Augener, 1906, pp. 188-191, pi. 7, figs. 145-147, pi. 8, figs. 148-152; M.C.Z. no. 2055-2060; West Indies, may be the same as P. diomedeae Benedict (1887, pp. 547-548, pi. 20, figs. 1-6) from Cape Hatteras and the Gulf of Mexico. The latter is incompletely described.
hartman: polychaetous annelids 21
Serpula nannoides Chamberlin, 1919b, p. 270, pi. 2, fig. 8; M.C.Z. no. 511; California.
[Serpula Columbiana :iohns,on] 1901, pp. 432-433, pi. 19, figs. 199-204; M.C.Z. no. 1896; Puget Sound to San Francisco, has long been con- sidered identical with Serpula vermicularis Linnaeus (see Chamberlin, 1919c, p. 269).
A REVISION OF THE SPECIES OF THE GENUS EUDISTYLIA BUSH
Type. Sabella vancouveri Kinberg (1867, p. 353; Vancouver Island).
Diagnosis. Paired branchial lobes equal, their ventral ends free and turned inward, or spirally coiled. The free dorsal ends of the branchial bases produced in a more or less flattened, simple or cleft, stiffened membrane (pi. 2, fig. 5).
Radioles numerous, usually simple, rarely divided, the stem approx- imately three-sided in cross section but the angles blunt. Pinnae arranged in double series on the inner sides of the radioles as character- istic of the SABELLINAE, without stylodes. Eye spots more or less numerous, rarely absent, arranged in a single or irregular double series on the outer sides of the radioles.
Collarette with dorsolateral notches in addition to the dorsal and ventral clefts. Oral membranes ruffled, thin, wide. Palpi lanceolate or foliaceous. Thorax with dorsal furrow and ventral plates. Thoracic notopodia with winged lanceolate setae and spatulate setae, the latter sharply set off; thoracic neuropodia with avicular uncini accompanied by pennoned setae. Abdomen with dorsal plates divided by a deep- longitudinal furrow. Abdominal parapodia with notopodial avicular uncini and neuropodial winged lanceolate setae.
Remarks. Dift"ers from Bispira Kroyer primarily in having spatulate setae instead of the scimitar-like setae in thoracic notopodia.
This genus is known only from the Pacific Ocean. Ten species have been assigned to it. These are listed below. Those considered syno- nyms are enclosed in brackets.
*[Eudistylia abbreviata Bush] 1904, p. 212; Alaska; is identical with E. vancouveri (Kinberg). The color was said to have been similar to that of E. gigantea, which is also identical with E. vancouveri. The collar is deep along the sides, and curves obliquely ventrally; setae and collar agree with those described for E. vancouveri (see Johansson, 1926, p. 14).
22 bulletin: museum of comparative zoology
Eudistylia brevicomata (Ehlers) Augener, 1926, p. 251 ; New Zealand, may be identical with Sabella ceratodaula Sehmarda Augener, 1926, p. 253 (see also Johansson, 1927, p. 138). Ehlers described the eye spots as disposed in two irregular rows. I have not seen this condition in any specimens of Eudistylia from the north Pacific.
Augener (1926, p. 253) has already discussed the possible identity of E. brevicomata Ehlers, »S'. ceratodaula Sehmarda and S. grandis Baird (1865, p. 160), all from New Zealand. It is likely that the descriptions of Sehmarda, Baird, Kinberg and Ehlers were all based on individuals which represent a single, widely distributed species. Such a conclusion cannot be made, however, without study of more collections from the areas in question.
It is of interest that Ehlers had at least one specimen of " Bispira polymorpha Johnson" from California, and that he observed no signif- icant differences between his own species and the specimen from John- son. Unfortunately it is not possible to know whether Johnson had sent him E. vancouveri or what is really E. polymorpha (see below). Probably it is the former, however, since the color pattern of it agrees with that of E. brevicomata. A pencilled note on the back of Johnson's label of M.C.Z. type no. 1941 reads, "1 sp. sent Ehlers Apr. 30, '03." Since it appears unquestionable that Johnson did base his description and illustrations on two species, and since his figure of the anterior portion of the animal (Johnson, 1901, fig. 179) agrees fairly well with the specimen remaining in the type jar, 1941, it is reasonable to assume that he had sent Ehlers the other species, E. vancouveri (see discussion under B. polymorpha, below).
'^Sabella ceratodaula Sehmarda 1861, from New Zealand, may be the same as E. brevicomata (Ehlers) 1905, from the same region, (see Augener, 1926, p. 248).
^[Eudistylia gigaiitca Bush] 1904, p. 210; is identical ■with.E. vancouveri (Kinberg). Johansson (1927, p. 135) has already indicated this syn- onymy. The radioles (according to Bush's description are marked with three conspicuous bands of maroon. The illustrations of Bush (1904, pi. 22, figs, a, c) however, show four bands of pigment. Johansson, on reexamining Kinberg's type of Sabella vancouveri, described four or five bands of pigment. In other respects, S. vancouveri and E. gigantea agree.
*[Eudistylia intermedia Bush] 1904, p. 214; California, is question- ably referred to E. polymorpha (Johnson) . Bush separated this species from E. polymorpha on the basis that its branchiae are more numerous, the eyes less conspicuous and the avicular uncini shaped somewhat
hartman: polychaetous annelids 23
differently. She may have had specimens of E. vancouveri (Kinberg) for comparison. Branchiae were described as being "brown or pinkish purple on the lower portion" with a "narrow, scarcely discernible pink band farther out." This agrees with E. polymorpha and not with E. vancouveri. The sjTionymy of E. intennedia and E. polymorpha is further indicated in that the eyes were "less conspicuous." The eyes of E. polymorpha are well marked, but less conspicuous because of the darker color of the radioles.
*[DisUjlia monterea Chamberlin] 1919b, p. 267; M.C.Z. no. 1941; California, belongs to Eudistylia, identical with E. polymorpha. The type specimen was originally one of Johnson's four types of Bispira polymorpha, the other tliree type jars (M.C.Z. nos. 1901, 1902, 1903) include E. vancouveri and two species of Schnizobranchia. That Johnson partly used type 1941 in his original description is certain, for he mentions the two color phases "purple or wTOe-color, and whitish or tawny," the former being what Chamberlin subsequently named D. vionterea, and the latter specimens of E. vancouveri (see also discussion under E. brevicomata and E. polymorpha) .
'^[Eudistylia plumosa Bush] 1904, p. 212; Alaska, is probably identi- cal with E. vancouveri (Kinberg). The thoracic avicular uncini re- semble those of Kinberg's species. E. plumosa is too incompletely known to permit certain identity. Nothing is known of the nature of the collar, the branchial lobes, the radioles, except that the collar has conspicuous ventral ends, and the branchiae are banded with pigment. Both of these characters agree, however, with E. vancouveri.
Bispira polymorpha Johnson, 1901, p. 428; M.C.Z. nos. 1901, 1902, 1903, 1941; Puget Sound to Pacific Grove, includes the following:
M.C.Z. no. 1901, from Port Orchard, is a Schizobranchia, probably S. nobilis Bush.
M.C.Z. no. 1902, from Puget Sound, includes two specimens, E. vancouveri (Kinberg) and Schizobranchia insignis Bush.
M.C.Z. no. 1903, from Humboldt Bay, is E. vancouveri (Kinberg)
M.C.Z. no. 1941, from Pacific Grove, includes a single specimen, hereby retained to represent the holotype of E. polymorpha (Johnson) .
M.C.Z. no. 1941 was used in the original description, as brought out above, under D. monterea. (See also E. brevicomata, above).
Question arises as to the status of the names, Bispira polymorpha Johnson (1901), Eudistylia intermedia Bush (1904) and Distylia monterea Chamberlin (1919). The first is clearly based on at least two species of Eudistylia, one the older E. vancouveri (Kinberg), the other
24 bulletin: museum of comparative zoology
what was subsequently made the type of Distylia monterea. Two procedures present themselves. (1) Reduce polymorpha to synonymy with vancouveri and retain monterea for the specimen M.C.Z. 1941, or (2) retain polymorpha for M.C.Z. 1941, reducing monterea to synonymy with it. The former procedure would be preferable in some respects were it not for the questionable status of E. intermedia. The existence of Bush's type is not known, and its identity with E. monterea not certain. Also, it predates Chamberlin's name, hence places monterea in the position of a possible synonym. Consequently I am retaining Johnson's type, M.C.Z. 1941, as the holotype of E. polymorpha and considering E. intermedia and D. monterea as synonyms.
Wudistylia tenella Bush, 1904, pp. 213-214; Alaska, resembles Demonax in (1) lacking eye spots, (2) having delicate pinnae, (3) the nature of the collar. It may be a specimen of the widely distributed Demonax leucaspius Kinberg.
Eudistylia vancouveri (Kinberg) 1865, p. 353; Vancouver Island, was redescribed by Johansson (1926, p. 13; 1927, p. 135) who had access to the type. It probably includes E. abbreviata Bush, E. gigantea Bush, E. plumosa Bush, and E. polymorpha, in part. Radioles have five to eight transverse pigment bands of alternating tawny and maroon. Each eye spot usually has an oblique splash of color. Collar has high dorsal lobes (pi. 2, fig. 2), deep dorsolateral clefts, oblique lateral lobes and broadly triangular ends (pi. 2, figs. 3 and 4). Dorsal branchial base is simple, not cleft (pi. 2, fig. 2). Dorsal oral membrane is provided with medial and lateral finger-like elongations (pi. 3, fig. 7).
Key to the accepted species of Eudistylia
1 . Branchial eye spots disposed in double rows E. brevicomata
1. Branchial eye spots in single rows 2
2. Dorsal side of branchial membrane deeply cleft, the two lobes thus formed
broadly overlapping (pi. 2, fig. 5); branchiae deep maroon except for a short band distally; ventral collar lobes not greatly produced and not
oblique E. polymorpha
2. Dorsal side of branchial membrane with rectangular flaps, not cleft (pi. 2, fig. 2); branchiae with numerous transverse bands of maroon and flesh-colored (preserved) ; ventral lobes of collar produced, oblique.
E. vancouveri
hartman: polychaetous annelids 25
A REVISION OF THE SPECIES OF THE GENUS PSEUDOPOTAMILLA BUSH
Type. Amphitrite reniformis Montagu.
Diagnosis. Paired branchial lobes equal, circular, not spiralled. The free dorsal ends of the branchial bases produced in a flattened, simple or cleft, membrane (pi. 2, fig. 8). Radioles numerous, simple, not divided; pinnae arranged in double series as characteristic of the SABEL- LINAE, without stylodes. Eye spots in single rows, disposed on the outer, lateral borders of the radioles.
Collarette with dorsal and ventral clefts, and dorsolateral incisions; otherwise as in Potaniilla Malmgren. Thoracic setae consist of lanceolate and spatulate setae in notopodia, and avicular hooks with companion pennoned setae in neuropodia. Abdominal setae include avicular hooks in the notopodia and lanceolate setae in neuropodia.
The species that have been assigned to Pscudopotainilla, and those assigned to other genera but which are of this genus, are listed below. SjTionyms are enclosed in brackets. New synonyms are preceded by an asterisk.
[*Pseudopotamilla anoculata Moore] 1905, pp. 564-566, pi. 37, figs. 23-27; Alaska, is a Megalomma, identical with M. splendida (Moore) new comb.
*[Pseudopotamilla brevibranchiata Moore] 1905, pp. 555-559, pi. 37, figs. 1-7; Alaska, is identical with Ps. occelata. These two were first separated on the observation that the latter was longer, had more elongated Ijranchiae, more conspicuous branchial eyes, and because of slight differences in the shapes of the avicular setae. Subsequent records of both species (Moore, Chamberlin, etc.) have shown that the branchiae are usually longer than first recorded for Ps. brevibranchiata and shorter than for Ps. occelata, and that the differences first noted, vary within wide ranges. Branchial eye spots vary, both as to size and number (see also Chamberlin, 1918, p. 179; Berkeley, 1930, p. 70).
IPseudopotamiUa debilis Bush, 1904, p. 204, pi. 36, figs. 23, 24, 26; California, cannot be identified with certainty. It may be either Ps. occelata Moore, or Ps. intermedia Moore.
[Pseudopotamilla debilis Moore] 1908, p. 360; Gulf of Georgia, de- posited in the U.S. National Museum, is a specimen of Mcgcdomma splendida (Moore) (see below).
Pseudopotamilla intermedia Moore, 1905, pp. 562-564, pi. 37, figs. 15-22; Alaska, probably includes Ps. reniformis Moore (1908, p. 359;
26 bulletin: museum of comparative zoology
Alaska, Berkeley, 1930, p. 70; British Columbia) and also Ps. macrops Chamberlin (1919a, p. 22; California). Characteristic features of this species are: (1) The dorsal branchial membrane is a narrow, rectangular flap, hardly overlapping (pi. 2, fig. 8) ; (2) the dorsalmost radiole has a slender tentacle, set some distance from the base of the pinnae (pi. 3, fig. 1) ; (3) the uncini in the last two or three thoracic segments are large, long-handled, in contrast to the smaller ones in more anterior segments (cf . pi. 3, figs. 4 and 5) ; (4) the dorsal collar lobes are free lappets, concealing the peristomium (pi. 2, fig. 8). Spatulate setae have a slender mucron (pi. 3, fig. 3).
*[Pseudopota7nilla lampra Chamberlin] 1919a, p. 22, is identical with Ps. occelata Moore. The cleft dorsal branchial membrane, collar, thoracic structures and branchiae are like those in Ps. occelata.
'^[Pseudopotamilla macrops Chamberlin] 1919a, p. 22, is identical with Ps. intermedia Moore. The type (M.C.Z. no. 177) is unique, con- sists of only head and three setigerous segments. The dorsal edge of the branchial membrane is a narrow, entire flap ; radioles have few eye spots, there are three on the left side, one on the right side; dorsal lobes of the collar are almost as high as the lateral lobes; the collar is as described for Ps. intermedia. The nature of the posterior thoracic segments cannot be ascertained.
Pseudopotamilla occelata Moore, 1905, pp. 559-562, pi. 37, figs. 8-14; Vancouver Island, is a widely distributed species in the northeast Pacific. It includes the following species, Ps. hrevihrancJiiata Moore, Ps. brevibranchia Chamberlin, Ps. lampra Chamberlin, Ps. paurops Chamberlin, Laonome oculifera Treadwell, and probably Ps. scotia Chamberlin. Characteristic features of this species are:
The branchial membrane has notched wings dorsally (pi. 2, fig. 6), dorsal lobes of the collar are low, exposing much of the peristomium; lateral lobes of the collar rise abruptly anterior to the collar fascicle; the lateral lobes are set off from the ventral lobes by a slight notch, the ventral lobes are continued along their ventralmost margins into two elongated lobes. Branchial eyes are black, conspicuous, rounded, or often with a faint short splash; ventral thoracic plates are rec- tangular, except the first which is shield-shaped (pi. 2, fig. 7).
Remarks. This species occurs commonly in the littoral zone, from Alaska south to southern California. It is often in clusters of tubes or colonies.
^[Pseudopotamilla panamaica Chamberlin] 1919b, p. 268; Panama, is a Megalomma, identical with M. wsiculosum (Montagu) cosmopoli- tan.
hartman: polychaetous annelids 27
*[Pseudopotamilla parva Chamberlin] 1919a, p. 21, is doubtfully re- ferred to Megalornma (see also p. 499).
*[Pseudopotamilla paurops Chamberlin] 1919a, p. 21; California, is identical with Pseudopotamilla occelata. The type is much like that of P. lampra.
Pseudopotamilla reniformis (Montagu) is a widely distributed, warm water species. Such records as have been reported from the northeast Pacific must be doubted, in view of the possible confusion of this species with P. intermedia Moore.
^[PseudopotamiUa scoiia Chamberlin] 1919a, p. 22; California, is doubtfully referred to P. occelata Moore. The type has been dried and is incomplete. It consists of the head and eight setigerous segments. The radioles are greatly shrunken and have lost much of the epithelial layer. The specimen agrees with P. occelata in having a notched dorsal branchial membrane, the dorsal collar lobes are low, the collar is straight, and the ventral tips are elongated.
^[Pseudopotamilla splendida Moore] 1905, pp. 564, pi. 37, figs. 23-27; iVlaska, is Megalornma splendida, new comb. Its terminal eyes are spiralled as in Branchiomma burrardum Berkeley (1930, p. 71). There are other reasons for regarding the two synon;yTxious, as is patent in comparing the descriptions of the two. Ps. anoculata Moore is also identical with M. splendida, as has been verified by examining the types in the U.S. National Museum.
""[Laonome oculifera Treadwell] 1914, pp. 222-223, pi. 12, figs. 39^3; California, is a Pseudopotamilla, identical with Ps. occelata Moore. Its type has been examined in the collections of the University of California, and found to agree with Moore's species.
Sabella (Potamilla) oligophthalmos Grube from the Philippines, is a Pseudopotamilla (see Augener, 1926, p. 253). It includes Potamilla lacinosa Ehlers, 1905, from New Zealand (see Augener, — 1926).
[Potamilla lacinosa Elilers] 1905, pp. 66-67, pi. 9, figs. 7-10; from New Zealand, is considered identical with P. oligophthalmos (Grube) from Singapore (see Augener, 1926, p. 253).
28 bulletin: museum of comparative zoology
BIBLIOGRAPHY
Arwidsson, I.
1907. Studien iiber die skandinavischen und arktischen Maldaniden nebst Zusammenstellung der i'lbrigen bisher bekannten Arten dieser Familie. Zool. Jahrb. Supp., 9:1-308, 12 pis.
AUGENER, H.
1923. Polychaeta. I. Polychaeten von den Auckland und Campbell- Inseln. Videns. Medd. Kjobenhavn, 75:1-115.
1906. Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mexico and the Caribbean Sea, and on the east coast of the United States, 1877 to 1880, by the U.S. Coast Survey Steamer "Blake." Westindische Polychaeten. Bull. Mus. Comp. Zool. Harvard, 43:91-196, 8 pis.
1922. tjber litorale Polychaeten von Westindien. Sitzber. Gesells. natf. Freunde, Berlin, 1922:38-53.
1925. Uber Westindische und einige andere Polychaeten-Typen von Grube (OErsted), Kr03'er, Morch und Schmarda. Publ. fra Univ. Zool. Museum, K0benhavn, 39:1-47, 3 figs.
1926. Polychaeten von Neuseeland. II. Sedentaria. Vidensk. Medd. nat. Forh. Kjobenhavn, 81:157-294, 22 figs.
Baird, W.
1863. Descriptions of several new species of worms belonging to the Annelida Errantia and Sedentaria or Tubicola of Milne Edwards. Proc. Zool. Soc, London, 1863:106-110.
1865. On new tubicolous annellides in the collection of the British Museum. Jour. Linn. Soc, London, 8:157-160, 1 pi.
Benedict, J.
1887. Description of ten species and one new genus of annelids from the dredgings of the U.S. Fish Comm. steamer "Albatross." Proc. U.S. Nat. Mus., 9:547-553, 6 pis.
Bergstrom, E.
1914. Zur Systematik des Polychaeten Familie der Phyllodociden. Zool. Bidr. Uppsala, 3:37-224, 81 figs.
Berkeley, E.
1923. Polychaetous annelids from the Nanaimo District. Part I. Syllidae to SigaHonidae. Contr. Canad. Biol. Fish., Toronto, 1:205-219, 1 pi.
1929. Polychaetous annelids from the Nanaimo district. Part 4. Chae- topteridae to Maldanidae. Contr. Canad. Biol. Fish., Toronto, 4:305-316, 1 pi.
1930. Polychaetous annelids from the Nanaimo district. Part 5. Am- mocharidae to Myzostomidae. Ibid., 6:65-77, 8 figs.
hartman: polychaetous annelids 29
Berkeley, E. and Berkeley, C.
1932. On a collection of littoral Polychaeta from the west coast of Van- couver Island. Ibid., 7:309-318.
1936. Notes on Polychaeta from the coast of western Canada. I. Spionidae. Ann. Mag. Nat. Hist., 18:468-476, 1 fig.
1938. Notes on Polychaeta from the coast of western Canada. II. Syllidae. Ann. Mag. Nat. Hist., ser. 11, 1:33-49, 12 figs.
Bush, K.
1904. Tubicolous annelids of the tribes Sabellides and Serpullides from the Pacific Ocean. Harriman Alaska Expedition, N. Y., 12:169-355, 24 pis.
Caullery, M.
1915. Sur les terebelliens du genre Pista Mgn. et en particulier sur les uncini de ces Annelides. Bull. Soc. ZooL, France, 40:68-78.
Chamberlin, R.
1918. Polychaetes from Monterey Bay. Proc. Biol. Soc. Washington,
31:173-180. 1919a. New polychaetous annelids from Laguna Beach, California. Jour.
Entom. ZooL, Pomona Coll., 11:1-23. 1919b. Pacific coast Polychaeta collected by Alexander Agassiz. Bull.
Mus. Comp. Zool., Harvard, 63:251-276, 3 pis. 1919c. The Annehda Polychaeta. Mem. Mus. Comp. ZooL, Harvard,
48:1-514, 80 pis. 1920. Report of the Canadian Arctic Expedition, 1913-18. Volume 9.
Annelids, Parasitic worms. Protozoans, etc. Part B. Polychaeta.
Ottawa, pp. 1-41B, 6 pis.
Ehlers, E.
1868. Die Borstenwiirmer nach systematischen und anatomischen Unter- suchungen dargestellt. Leipzig. 748 pp., 24 pis.
1879. Reports on the results of dredging under the supervision of Alex- ander Agassiz, in the Gulf of Mexico, by the United States coast survey Steamer "Blake," Lieutenant-Commander C. D. Sigsbee, U.S.N. Commanding. Preliminary report on the worms. Bull. Mus. Comp. ZooL, Harvard, 5:269-274.
1887. Report on the annelids of the dredging expedition of the U.S. coast survey Steamer "Blake." Mem. Mus. Harvard, 15:335 pp., 60 pis.
1905. Neuseelandische Anneliden. Abh. K. Gesells. Wiss. Gottingen, Math.-Phys. Kl., 3:1-80, 9 pis.
Essenberg, C.
1917. On some new species of Aphroditidae from the coast of California. Univ. Calif. Publ. ZooL, Berkeley, 16:401-430, 7 pis.
Fauvel, p.
1923. Polychetes errantes. Faune de France, Paris, 5:1-488, 188 figs.
30 bulletin: museum of comparative zoology
Gravier, C.
1901. Contribution a I'etude des Ann^lides Polychetes de la Mer Rouge. Nouv. Arch. Mus., Paris, (4)3:147-268, figs. 160-285, 4 pis.
Grube, E.
1860. Beschreibung neuer oder wenig bekannter Anneliden. Arch. Naturg., 26.1:71-118, 3 pis.
Hartman, O.
1938a. Descriptions of new species and new generic records of polychaetous
annelids from California of the families Glyceridae, Eunicidae,
Stauronereidae and Opheliidae. Univ. Calif. Publ. Zool., 43: 93-112,
63 figs. 1938b. Review of the annelid worms of the family Nephtyidae from the
northeast Pacific, with descriptions of five new species. Proc. U.S.
Nat. Mus., 85:143-158, 6 figs.
Hessle, C.
1917. Zur Kenntnis der terebellomorphen Polychaeten. Zool. Bidr. Upp- sala, 3:39-258, 5 pis., 66 figs.
Johansson, K.
1926. Bemerkungen iiber die Kinbergischen Arten der Familien Her- mellidae und Sabellidae. Ark. Zool., Stockholm, 18 A.7:l-28, 9 figs.
1927. Beitrage zur Kenntnis der Polychaeten Familien Hermellidae, Sabellidae und Serpulidae. Zool. Bidr. Uppsala, 11:1-184, 15 figs., 5 pis.
Johnson, H.
1897. A preliminary account of the marine annelids of the Pacific coast, with descriptions of new species. Part 1. Euphrosynidae, Am- phinomidae, Palmyridae, Polynoidae and Sigalionidae. Proc. Calif. Acad. Sci. Zool., 1:153-190, 6 pis.
1901. The Polychaeta of the Puget Sound region. Proc. Boston Soc. Nat. Hist., 29:381-437, 19 pis.
Langerhans, p.
1879. Die Wurmfauna von Madeira. Zeits. wiss. Zool., 33:271-316, 5 pis.
McIntosh, W.
1 885 . Report on the Annelida Polychaeta collected by H . M . S. 'Challenger" during the years 1873-76. Challenger Reports, 12:1-554, 55 pis.
Mayer, A.
1900. An Atlantic "Palolo" Staurocephalus gregarious. Bull. Mus. Harvard, 36:1-14, 3 pis.
1902. The Atlantic Palolo (Eunice fucata). Bull. Brooklyn Mus., 1:93-103, Ipl.
hartman: polychaetous annelids 31
Monro, C. C. A.
1928. Polychaeta of the families Polynoidae and Acoetidae from the vicinity of the Panama Canal, collected by Dr. C. Crossland and Dr. Th. Mortensen. Jour. Linn. Soc. London, 36:553-576, 30 figs.
1930. Polychaete worms. Discovery Reports, 2:1-222, 91 figs.
1933a. The Polychaeta Errantia collected by Dr. C. Crossland at Colon in the Panama region and the Galapagos Islands during the Expedition of the S.Y. "St. George." Proc. Zool. Soc, London, 1933:1-96, 36 figs.
1933b. The Polychaeta Sedentaria collected by Dr. C. Crossland at Colon in the Panama Region, and the Galapagos Islands during the expedi- tion of the S.F. "St. George." Ibid., 1933:1039-1092, 31 figs.
Moore, J.
1911. The polychaetous annelids dredged by the LT.S.S. "Albatross" off the coast of southern California, in 1904. Euphrosinidae to Goni- adidae. Proc. Acad. nat. soi., Philadelphia, 1911:234-318, 7 pis.
1923. The polychaetous annelids dredged by the U.S.S. "Albatross" off the coast of southern California in 1904. Spionidae to Sabellariidae. Ibid., 1923:179-259, 2 pis.
POURTALES, L. F. DE
1869. Contributions to the fauna of the Gulf Stream at great depths. Bull. Mus. Comp. Zool. Harvard, 1:103-120.
QUATREFAGES, A. DE
1865. Histoire naturelle des Anneles marins et d'eau douce. Annelides et Gephyriens. 3 vols. (Paris, Libr. Encycl. de Roret).
Schmarda, L.
1 86 1 . Neue wirbellose Thiere beobachtet und gesammelt auf einer Reise um die Erde, 1853 bis 1857. 164 pp., 22 pis.
Seidler, H.
1924. Beitrage zur Kenntnis der Polynoiden. Arch. Naturg., 89, (Abt. A., Heft), pp. 1-217, 22 figs.
Treadwell, a.
1914. Polychaetous annelids of the Pacific coast in the collection of the zoological museum of the University of California. Univ. Calif. Publ. Zool., 13:175-234, 2 pis.
Webster, H.
1884. Annelida from Bermuda, collected by G. Brown Goode. Bull. U.S. Nat. Mus., no. 25, pp. 305-327, 6 pis.
EXPLANATION OF PLATES
PLATE 1
HaRTMAN POLYCHAETOUS AnNBLIDS
PLATE 1.
Fig. 1. Nephtys bucera Ehlers. Foot from eleventh posteriomost segment,
in anterior view, x 22. Fig. 2. Nephtys discors Ehlers. Foot from posterior fourth of body, in
anterior view, x 12. Fig. 3. Same. Foot from anterior fourth of body, in anterior view, x 12. Fig. 4. Scoloplos acnieceps Chamberlin. Parapodium from thirtieth seg- ment, X 35. Fig. 5. Myxicola infundibulum (Renier). Anterior end in ventral view, of
specimen from Cahfornia, x 11.
Same. Anal region, showing distribution of pigmented spots, x 11.
Same. Anterior end in dorsal view, first setal fascicle indicated, x 11.
Same. Larger abdominal uncinus, x 700.
Same. Smaller abdominal uncinus from same foot as that in fig. 8,
and alternating with it, x 700.
Same. Anterior end in lateral view, setae omitted, x 11.
Same. Four anterior abdominal segments in lateral view, showing
distribution of uncini (smaller dots) and ocular points with respect
to setal fascicles, x 1 1 .
|
Fig. |
6. |
|
Fig. |
7. |
|
Fig. |
8. |
|
Fig. |
9. |
|
Fig. |
10. |
|
Fig. |
11. |
BULL. MUS. COMP. ZOOL.
Hartman. Polychaetous Annelids. Plate 1.
PLATE 2
HaRTMAN POLYCHAETOUS ANNELIDS
PLATE 2.
Myxicola infundibulum (Renier). Thoracic hook, x 700. Eudistylia vancouveri (Kinberg). Collar and base of branchial membrane, in dorsal view, x 11.
Same. Anterior end, branchiae omitted, in left lateral view, x 11. Same. Ventral ends of collar, from specimen killed out of tube, x 1 1 . Eudistylia poly mar pha (Johnson). Collar and base of branchial membrane, in dorsal view, x 11.
Pseudopotamilla occelata Moore. Anterior end, showing notched branchial base and low dorsal collar lobes, x 11.
Same. First four thoracic ventral plates, from specimen killed out of tube, x 11. Fig. 8. Pseudopotamilla intermedia Moore. Anterior end, showing entire branchial base and well rounded dorsal collar lobes, x 11.
|
Fig. Fig. |
1. 2. |
|
Fig. Fig. Fig. |
3. 4. 5. |
|
Fig. |
6. |
|
Fig. |
7. |
BULL. MUS. COMP. ZOOL.
Hartman. Polychaetous Annelids. Plate 2.
PLATE 3
HaRTMAN POLYCHAETOUS ANNELIDS
PLATE 3.
Fig. 1. Pseudopotamilla intermedia Moore. Dorsalmost radicle from right
side, showing tentacular membrane and rectangular branchial base
at left, X 11. Fig. 2. Same. Ventralmost radiole from right side with attached palpus,
from same specimen as fi.g. 1 , x 11. Fig. 3. Same. Spatulate seta from first thoracic segment, showing tiny
mucron, x 347. Fig. 4. Same. Uncinus and companion pennoned seta from last thoracic
segment, x 347. Fig. 5. Same. Uncinus and companion pennoned seta from second last
thoracic segment, from same specimen as that of fig. 4, x 347. Fig. 6. Same. Neuropodial setae from second abdominal segment, x 347. Fig. 7. Eudisfylia vancouveri (Kinberg). Base of dorsalmost radiole from
right side, showing tentacular membrane and rectangular branchial
base at left, x 11. Fig. 8. Sabellaria nanella Chamberlin. Palea from inner series of opercular
crown, X 45. Fig. 9. Same. Palea from middle series of opercular crown, x 45. Fig. 10. Same. Palea from outer series of opercular crown, showing simple,
smooth distal spine, x 45.
BULL. MUS. COMP. ZOOL.
Hartman. Polychaetous Annelids. Plate 3.
Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. LXXXV, No. 2
SOME PARASITIC WORMS IN THE HELMINTHOLOGICAL
COLLECTION OF THE MUSEUM OF
COMPARATIVE ZOOLOGY
By J. H. Sandground
CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
December, 1938
PUBLICATIONS OF THE
MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD COLLEGE
There have been published of the Bulletin, Vols. I to LXXXII, and LXXXV, No. 1 & 2; of the Memoirs, Vol. I to LIV, No. 1, 2 & 3, and Vol. LV.
The Bulletin and Memoirs are devoted to the publication of original work by the Officers of the Museum, of investigations carried on by students and others in the different Laboratories of Natural History, and of work by specialists based upon the Museum Collections and Exploration.
These publications are issued in numbers at irregular intervals. Each number of the Bulletin and of the Memoirs is sold separately. A price list of the publications of the Museum will be sent on application to the Director of the Museum of Comparative Zoology, Cambridge, Massachusetts.
Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. LXXXV, No. 2
SOME PARASITIC WORMS IN THE HELMINTHOLOGICAL
COLLECTION OF THE MUSEUM OF
COMPARATIVE ZOOLOGY
By J. H. Sandground
CAMBRIDGE, MASS., U.S.A.
PRINTED FOR THE MUSEUM
December, 1938
No. 2. — Some Parasitic Worms in the Helniinthological Collection of the Museum of Comparative Zoology
By J. H. Sandground
I. Descriptions of some parasitic worms from Solenodon paradoxus Brandt (Insectivora)
Among the rarest of extant mammals are the two species of Soleno- don, S. paradoxus Brandt, 1833 and S. cubanus Peters, 1861, which together constitute the very distinctive insectivore family Soleno- dontidae. Nocturnal in their habits, these fairly large, shrew-like animals live in subterranean galleries or in hollow trees and are restricted in distribution to heavily wooded mountains or rain forests on two islands in the Caribbean Sea. Although the small series of study specimens that it has been possible for a few of the world's largest museums to assemble indicates that these animals have always been difficult to procure, it is now generally supposed that with the development of agriculture in Cuba, the species S. cubanus has become extinct. Furthermore, since so few specimens of S. paradoxus have been secured on the island of Haiti, it was feared that this species had also suffered a similar fate. That this fear was not entirely warranted was proved by a German collector who, in 1935, succeeded in cap- turing 28 specimens in San Domingo.
Desiring specimens for particular anatomic studies. Professor George B. Wislocki of the Harvard Medical School commissioned the services of Mr. William J. Clench of the Museum of Comparative Zoology, who in the summer of 1937 was leading an expedition into the interior of the Republic of San Domingo. With the aid of native collectors, Mr. Clench was eventually successful in obtaining 3 adult specimens of S. paradoxus. Subsequently, while working on the formalin preserved body of one of these animals. Dr. Wislocki ob- served that the subperitoneal muscles of the pelvis were riddled with encysted parasites. These he kindly turned over to the writer, together with the viscera of this and another specimen. The muscle-invading parasites were found to be the larvae of an as yet unidentified thorny- headed worm (Acanthocephala) which I also found in large numbers attached to the mesenteries and diaphragm and encysted under the capsule of the liver and on the pericardium. Search of the intestine revealed numerous specimens of tliree other classes of parasitic worms : roundworms in the duodenum, masses of tapeworms in the lower part of the small intestine, and flukes in the colon.
36 bulletin: museum of comparative zoology
In view of the rarity of the host it is not surprising that the literature of helminthology mentions no previous examination of Solenodon. Our studies indicate that at least two of the parasites belong to hitherto undescribed species. On the other hand, the trematode belongs to a genus whose many species are so similar in morphology, that I deem it undesirable to distinguish it with a new specific name.
The type material of the new species described in the following pages is deposited in the helminthological collection of the Museum of Comparative Zoology at Cambridge.
CESTODA
Hymenolepis wislockii spec. nov.
Numerous specimens of this tapeworm were found in the lower part of the small intestine. Small developmental forms ranging upwards in length from 4 mm. and consisting of a few growing proglot- tids and a scolex were plentiful in the upper jejunum and duodenum. The longest strobilae composed of several hundreds of narrow segments measure in the preserved state up to 80 mm. in length, with an average maximum width of 0.95 mm. Much shorter specimens are numerous. The scolex (fig. 1) presents a variable appearance according to the state of muscular contraction and the position of the rostellum which when everted from its spacious pouch forms a columnar protuberance 0.17 mm. in length and 0.15 mm. wide. Crowning the rostellum is a ring of from 38 to 40 hooks which measure 25 ^t to 28 /x from the pointed iblade tip to the base of the handle. The guard, about 9/x long, pro- jects almost parallel to the blade. Figure 2 depicts two hooks that were teased apart from the rostellum and manoeuvred into a single optical plane for camera lucida drawing. When the hooks remain in situ they may present a very different appearance and, depending upon the angle of vision, a wholly erroneous conception as to their size. Scolex about 0.35 mm. wide, with suckers from 0.11 to 0.13 mm. in diameter partially covered by a cuticular fold. Musculature well developed, there being 20 to 22 longitudinal bundles consisting of 7 to 10 muscle fibers in addition to the transverse muscle fibers. Ventral and dorsal excretory vessels pass ventral to the nerve cord and genital ducts. In immature segments the three developing testes lie in a transverse line; in more mature segments (fig. 3) the median testis is pushed forward on the aporal side. Cirrus sac measures 0.17 mm. by 0.06 mm. In addition to the seminal vesical coiled within the cirrus sac there is a voluminous external seminal vesical which in mature
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segments extends to the middle of the segment and is persistent even in gravid egg-laden segments. Cirrus, devoid of spines, opens into a scarcely perceptible genital atrium near the middle of the segment margin. Ovary small, slightly lobed, situated mesially together with a very inconspicuous vitelline gland. There is an ampulliform dilation
Fig. 1. Hymenolepis wislockii Sandground. Fig. 2. Hymenolepis wislockii Sandground. Fig. 3. Hymenolepis wislockii Sandground.
Scolex with extruded rostellum. Isolated booklets. Mature segment.
of the oviduct or receptaculum seminis situated ventral to the external seminal vesical. The vagina runs close to the posteroventral edge of the cirrus sac to open by the side of the cirrus. The uterus when fully formed spreads throughout the segment. The ova in utero in balsam mounts measure from 38 to 41 ju by 30 to 35 /x, but as found extruded naturally from the segment into the contents of the host's gut they measure 60 to 64 ju by 46 to 50 m in outside diameter.
38 bulletin: museum of comparative zoology
Species Relations
The genus Hymcnolepis Weinland, 1858, is perhaps the largest in the entire field of helminthology, some 300 species parasitic in a wide variety of birds and mammals being attributed to it. In an effort to facilitate identification, Mayhew (1925) suggested a segregation of the members of this unwieldy group into 3 genera, but for reasons that have been given by several authorities (e.g. Fuhrmann 1932) these genera have had to be suppressed. In view of the magnitude of the genus and the difficulty of distinguishing many of its species on the basis of descriptions, it is common practise among present day workers to establish the taxonomic status of members by comparing it with those species that have been described from phylogenetically related hosts. The validity of this procedure depends upon the assumption of an ecologically, if not physiologically, determined specificity of the parasite for its host. Obvious as are the potential difficulties in adopting a procedure whose reliability is still largely theoretical, the exigencies of the moment are such that the practise cannot be aban- doned, and consequently I shall restrict the discussion of the taxonomic status of the species described here to a comparison with the 30 species of Hymenolepis which have already been recorded from Insectivora. The names of these species together with the morphological features that are deemed of prime difi"erential value are tabulated below.
In view of the plasticity of the body of tapeworms and the changes in topographic relations that the gonads may assume at different stages in growth as well as in death (depending upon fixation methods), the number and shape of the rostellar hooks has been found to provide the most reliable means of distinguishing between species of Hymeno- lepis. On this basis the species here described shows greatest resem- blance to H. bacillaris (Goeze, 1782) and H. tiara (Dujardin, 1845) ^ The present species, however, differs from both of the aforenamed species in its much larger cirrus pouch and the size of the eggs as well as in other features. It is consequently believed that the species is new, and it is named H. wislockii as a token of my appreciation to the donor of the material.
' In this connection it may be noted that while H. tiara is stated by Baer (1928) to have 38 to 40 hooks, it is also described (Baer 1932) to have only 30 to 32 hooks. Whether this difference is an index of intraspecific variation or indicates the existence of two species remains to be shown.
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40 bulletin: museum of comparative zoology
TREMATODA
Brachylaemus sp.
From the colon of one of the two specimens of Solenodon paradoxus dissected, eleven greyish colored flukes were recovered. These small worms in the preserved condition are almost round in cross section; the two bluntly attenuated extremities tend to bend ventrally so that from the side the worms are crescentic in shape. The pressure used in flattening specimens for the preparation of toto mounts causes a slight increase in width and may also derange other proportions of the body. The following measurements were all taken from toto mounts (fig. 4). Length 2.2 to 2.85 mm. Breadth at widest point 0.65 to 0.75 mm. Cuticle over preacetabular zone ornamented with minute spines. The spinosity is more difficult to detect in some individuals than in others. Oral sucker circular to elliptical in outline with diameter ranging from 0.25 to 0.35 mm. The mouth opening is partially covered by a fold of cuticle. Pharynx more or less spherical, 0.16 to 0.18 mm. in cross section. Intestinal crura bend anteriorly around the pharynx and subsequently run a very sinuous course, partially covered by the uterine coils, practically to the caudal extremity. Acetabulum, at junction of anterior third of body, 0.27 to 0.31 mm. in diameter. The short main stem of the excretory vesical opens subterminally on the ventral surface. Vitellaria, consisting of a narrow band of follicles, extend from the posterior level of the acetabulum to the level of the anterior testes; the transverse vitelline ducts dilate to form a con- spicuous reservoir crossing the ovary. Unsatisfactory fixation and the thickness of our specimens make for poor differential staining so that for the determination of the relationship of the genital ducts it has been necessary to rely largely upon serial sections. The gonads lie tandem in the median axis in the posterior quarter of the body with the ovary situated between and practically contiguous with the testes. The testes and ovary are all regular in outline; the latter organ (0.18 to 0.25 mm. in diameter) is only slightly smaller than the two testes. Even though Odhner's definition of the Brachylaemidae mentions the absence of a receptaculum seminis, our sections reveal a thin-walled sperm-containing dilation of the complex of ovarian ducts, just below the ovary. This we can only interpret as a seminal re- ceptacle. Baer (1928), Werby (1925), and others have also depicted a receptacula seminalis in various species of this genus, but Joyeux et al (1934) deny its presence, interpreting the structure as an elongate ootype. No trace of Laurer's canal was seen. The uterus is very
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41
lirm.
Fig. 4. Brachylaemus sp. Ventral view of toto-mount.
Abbreviations: o.s.-oral sucker, p. -pharynx, a-acetabulum, u-uterus, v-vitellaria, c-cirrus organ, m-metraterm, ti and t2-testes., o-ovary, i-in- testinal caeca, e.p.-excretory pore.
42 bulletin: museum of comparative zoology
voluminous, its coils (containing countless golden color eggs which measure 13-16^ by 24-28)u) extend beyond the acetabulum to the region of the pharynx. The metraterm and the cirrus pouch open into a common genital atrium, which is situated in the median ventral line well in the zone of the anterior testis. In toto mounts the genital pore is placed over the center of the testes; in lateral view it opens ventrally at the side of the testes. "
Systematic Relations
Members of the genus Brachylaemus (syn. Harmostomum) are very muscular, and in life the body is capable of considerable extension, more especially in the anterior region. They lose their motility soon after the death of their host and the body contracts markedly, some- times producing superficial wrinkling. Not only do measurements become a poor criteria of the species as a result of variations in con- traction, but the intestinal caeca take on undulations and the position of the genital pore relative to the anterior testis may be altered. The morphology of the organs varies apparently also with the state of maturity. As a consequence of these variations, the identification of species becomes a very difficult matter as attested by Dollfus (1934 and 1935) and by Joyeux, Baer and Timon-David (1934), who have recently assayed a revision of the genus, whose representatives are found in several orders of birds as well as in 3 orders of mammals.
The names of eleven species parasitizing members of the Insec- tivora are listed in the key catalogue of Stiles and Stanley (1932). Although many of these have been placed in synonymy by Baer (1928) and by Dollfus (1934), considerable uncertainty still exists regarding the validity of the remaining species. While it is to be hoped that life history studies will eventually elucidate the taxonomic status of many, the demonstrated fact that at least some species do not exliibit a high order of host specificity will not tend to reduce the existing uncertainty (see KruU, 1933 and Dollfus, Callot and Desportes 1935). Rather than complicate the already perplexing problem I think it desirable to ascribe no specific name to the species here described. It may however be noted that the species does not appear to differ significantly from B. helicis (Meckel, 1846) as recently described by Baer (1928) from European hedgehogs, nor from the species B. opisthotrias (Lutz, 1895) from South and North American opossums, as described recently by Dickerson (1930).
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NEMATODA
Shattuckius shattucki gen. et spec, no v.
Slender worms disposed to be loosely coiled in two or three whorls. Cuticle very faintly striated transversely and more conspicuously marked by about 16 longitudinal lines or elevated crests. There is an asjTnmetric inflation of the cuticle of the head for a distance of about 0.075 mm. (fig. 5). Cervical papillae apparently absent. Buccal
Fig. 5. Shattuckius shattucki Sandground. Anterior extremity.
cavity vestigial; without obvious lips but surrounded by 4 cephalic papillae. Oesophagus claviform, 0.31 mm. and 0.34 mm. long in male and female respectively; encircled by nerve ring near its middle.
Male 3.3 to 3.5 mni. long and 0.075 mm. maximum width. Female 5.4 to 5.7 mm. long with 0.085 mm. maximum width. Tail of female (fig. 6) 0.11 to 0.12 mm. long with a spinous process at its bluntly rounded tip. Vulva located 1.3 to 1.4 mm. from the caudal extremity; the cuticle around its rim (fig. 7) is slightly inflated making the vulva fairly conspicuous. Ovejectors about 0.16 mm. long. Eggs with very thin shells measuring in utero 48 to 50 ju by 32-35 m-
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Bursa of male (fig. 8) relatively spacious. There is a distinct though small dorsal lobe. Lateral lobes adorned with a punctate striation
Fig. 6. Shattuckius shattucki Sandground. Tail extremity of female.
along their internal postero-lateral margins. There is a small accessory membranous lobe, apparently devoid of supporting ribs. Prebursal papillae and telamon apparently absent. The long, relatively slender
Fig. 7. Shattuckius shattucki Sandground. Region of vulva.
externo-dorsal ray originates from a common trunk with the robust dorsal ray and is the only ray that does not extend to the margin of the bursa. The dorsal ray gives off serpentine lateral rays before
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bifurcating. On each branch of the divided dorsal ray, at about the mid-point there is an external bud, representing a vestigial digital process. Lateral rays about equal in size and well separated from each other. Ventral rays subequal, divergent from their common trunk to the bursal margin. Spicules (fig. 9), equal in size, about TO/z long, sparsely punctate on their surface. They are of unusually simple form,
Fig. 8. Shattuckius shatiucki Sandground. Posterior extremity of male.
devoid of lateral angular prominences and having only a single, very faint crest running longitudinally. It requires the highest possible magnifications to show that the spicule tips do not terminate in simple, acute points, but that they are cleft for a short distance to present two or possibly three splinter-like processes united by a transparent membrane. There is a very faintly cuticularized guber- naculum about 25 /x in length. Systematic Relations. From Stiles and Stanley's Key Catalogue (1932) we learn that the only representatives
46
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of the Trichostrongylidae recorded from the Insectivora are a number of species of Viannaia (HeHgmosominae) . The parasite described above, therefore, appears to be the first didelphous form, or member of the subfamily TrichostrongyHnae, to be reported from this order of mammals. A review of the literature indicates that our species is clearly a new form, but, because the number of genera in the Tricho- strongyHnae has been multiplied to such an extent by recent workers, the establishment of its taxonomic affinities has not proved simple. Superficially the species appears to be related to Cooperia, but the spinous process that tips the tail of the female and the subequality in size of the ventro-ventral and I atero- ventral rays as well as other
1^
Fig. 9. Shattuckius shattucki Sandground. Spicule isolated by dissection.
characters, immediately separate it from members of this genus. In respect to these same features it appears to show greater resemblance to the genera Molineus Cameron, 1923 and Maciella Travassos, 1935. From both of these genera, however, it difl^ers in the division of its dorsal ray and in the relatively plain structure of its spicules. We should prefer to regard characters of this order as having no more than specific value, in this way avoiding the oftentimes dubious creation of new genera, but the taxonomic standards that have been so generally adopted leave no alternative but to create a new genus which I name in homage to my colleague Dr. George C. Shattuck and briefly define as follows :
TrichostrongyHnae. Cuticle faintly striated transversely, and with about 16 longitudinal crests. With asymmetrical cephalic inflation
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but without cervical alae or cervical papillae. Vestigial buccal capsule. Female with mucronate tail; vulva postmedian. Eggs segment in utero. Bursa ample, with distinct dorsal lobe. Internal surface of lateral lobes ornamented along posterior borders. A very small accessory membrane present. Ventro-ventral and latero-ventral rays subequal and extending to bursa margin. Lateral rays about equally
Fig. 10. Shattuckius shattucki Sandground. Lateral lobe of the bursa with supporting rays.
long and well separated. External-dorsal, long and slender, but not at- taining margin, arises from common trunk with the massive dorsal ray. Dorsal gives off lateral branches before bifurcating into two halves, each of which has a small aborted process near its middle. Spicules relatively regular in outline, with well developed knob cephalad and cleft terminally into two or three parallel splinters. Gubernaculum acicular, very poorly chitinized. Prebursal papillae apparently ab- sent.
Genotype. S. shattucki.
Habitat. Duodenum of Solenodon paradoxus, Santo Domingo.
48 bulletin: museum of comparative zoology
BIBLIOGRAPHY
Baer, J. G.
1925. Sur quelques cestodes du Congo Beige. Rev. Suisse de Zool., 32,
p. 239. 1928. Contribution a la faune helminthologique de Suisse. Ibid., 35, p. 27. 1932. Contribution a la faune helminthologique de Suisse. Ibid., 39, p. 1.
DiCKERSON, L. M,
1930. A new variety of Harmostomum opisthotrias from the North American opossum, Didelphys tnrginiana, with a discussion of its possible bearing on the origin of its host. Parasitology, 22, p. 37.
DoLLFus, R. Ph.
1934. Sur quelques Brachylaemas de la faune Francaise recolt^s princi- plement a Richelieu (Indre et Loire) I. Ann de Parasit., 12, p. 551 II. Ibid. 13, p. 52
DoLFus, R. Ph., Callot, J. et Desportes, C.
1935. Infestation experimentale de Strigiformes par un Brachylaemus, Ibid., 13, p. 12.
FUHRMANN, O.
1932. Les T^nias des Oiseaux. M^moires de I'Univ. de Neufchatel. 8, pp. 382.
JoYEUx, Ch., Baer, J. G. et Timon-David, J.
1934. Recherches sur les Trematodes du genre Brachylaemus Dujardin. Bull. Biol. France Belg., 68, p. 385.
Krull, W. H.
1933. The opossum, Didelphys virginiana Kerr, a new host for Brachy- laemus spinulosum (Hofmann, 1899). Proc. Helminth. Soc. Wash- ington, 2, p. 98.
Mahew, R. L.
1925. Studies on the avian species of the cestode family Hymenolepidae. Illinois Biol. Monographs, 10, No. 1.
Stiles, C. W. and Stanley, S. F. <• 1932. Key catalogue of parasites reported for Insectivora. Bull. No. 159, National Institute of Health, Washington, D.C.
Werby, H. J.
1928. On the trematode genus Harmostomum, with description of a new species. Trans. Amer. Micros. Soc, 47, p. 68.
sandground: parasitic worms 49
2. A redescription of Tetrapetalonema digitata (Chandler, 1929) comb, no v., a filariid parasite of Gibbon Apes, with an enum- eration of its congeners.
Under the designation Dirofilaria (?) digitata, n. sp., Chandler (1929) described three female filariid nematodes found in the ab- dominal cavity of a Hylobates hoolock, which had died in the Calcutta Zoological Garden. In 1933, the present writer recorded the occurrence of the same species in an Annamese gibbon, Hylobates leucogenys from French Indo-China, but again because of the absence of male worms it was not possible to erase the mark of interrogation originally inserted by the discoverer of the species to emphasize the element of un- certainty in his generic designation.
Recently the Museum received a collection of parasitic worms that, in an incidental way, had been secured by the Asiatic Primate Expe- dition under the leadership of Harold J. Coolidge, Jr. In the collection were specimens of Dirofilaria pongoi Vogel and Vogelsang, 1930, taken from (1) an orang-utan, Pongo pygmaeus and (2) Hylobates funereus^ at Abai, in N. Borneo. Also there were several bottles containing very large numbers of Chandler's above-mentioned species which forms the subject of the present paper. This parasite had been found in a majority of adult gibbons, Hylobates lar entelloides (Geoffroy) taken at an altitude of 1000 m. on Mt. Intanon, near Cheingmai in northern Siam. At lower altitudes and in other portions of their range, the gibbons appeared to be free of filariasis. The collectors were greatly impressed by this epidemiological observation, and were also sur- prised that no signs of disease was manifested by apes so heavily in- fected that the peritoneal cavity and sometimes also the pleural cavity appeared to be crammed, "like a bowl of vermicelli" with crawling masses of worms. ^
A study of this abundant material now enables us to describe for the first time the male of Chandler's D. digitata and to transfer the species to the genus Tetrapetalonema.
' This appears to be the first record of D. pongoi in a gibbon.
' From the field notes of Professor A. H. Schultz of Johns Hopkins Medical School, and Dr. C. R. Carpenter of Bard College, two members of the expedition, we learn that infanta (i.e. under one year of age) were rarely infected; 12 of 19 juvenile gibbons carried sparse in- fections, while only two out of eighty adults were apparently free of filariid worms. In two of the latter it was noted that the ovaries were cystically enlarged with worms penetrating into the substances of the organ. In contrast with the high incidence and great intensity of infection with T. digitata that was encountered in gibbons captured in the forests, it was found that families of the same host species that lived segregated outside of the forest were free of infection. It seems possible that this observation may have special significance from the standpoint of the probable insect vector of the parasite.
50 bulletin: museum of comparative zoology
We desire here to express our appreciation to Dr. O. E. McCoy of the University of Rochester, New York, who presented to the Hel- minthological Collection of our Museum valuable specimens of T. marmosetae Faust, 1935, T. atalensis McCoy, 1936, and T. parva McCoy, the three species at present embraced in the genus. The examination of this material has been of great aid in considering the taxonomic status of the forms involved.
Tetrapatelonema digitata (Chandler, 1929)
The following account is based on a study of more than twenty representatives of each of the sexes taken from several specimens of the host.
Conforming with its congeners, this species is a very slender, thread- like worm. A marked attenuation of the body commences at an ap- preciable distance from the extremities. The flagelliform nature of the head and tail ends thus produced is more conspicuous in this species than in those described from New World monkeys. There is no noticeable guttural dilatation or annular swellings in other parts of the body such as are characteristically found in T. marmosetae and T. atalensis (cf. McCoy, 1936).
Although the cuticle in this genus has invariably been described as smooth or unstriated, close examination of specimens of probably all species which have not been subjected to complete clearing will show that the cuticle is striated with very fine lines at about 1.5^l intervals.
The anterior end of the body is bluntly rounded and about 30 )U wide in T. digitata. There is no semblance of a reinforced integumentary shield, cuticularized cap or analogous cephalic ornamentation such as is found in some of the genera that have incorporated in the sub- family Setariinae of Yorke and Maplestone. In a frontal view of the head (fig. 11) as well as in lateral view (fig. 12), eight delicate cephalic papillae in the subdorsal and subventral positions can be made out. They are arranged in a manner common among genera in theFilariidae. Although in frontal view the papillae may appear to be almost con- tiguous, they are actually well separated and it would be misleading to describe them as "twinned papillae at each corner of a rectangular peribuccal plate," whereby the inference could be drawn that, by a process of approximation or fusion, the papillae have been doubled, in analogy with the "double papillae" frequently seen on the lips of ascarids.
The lateral papillae, or, more correctly, amphids, are in the form of
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minute slit-like organs which are rather difficult to discern on the sur- face of the cuticle. These organs have by error been described by both
Fig. 11. Tetrapetalonema digitata. Frontal view of head.
Faust and McCoy as located in the mid-dorsal and mid-ventral fields. The minute oral aperture is simply a rounded pit without an ele- vated rim. It is difficult to get a clear impression of the structure of the
Fig. 12. Tetrapetalonema digitata. Anterior extremity in lateral view.
alimentary canal immediately behind the mouth owing to the region being obscured by strands of nerve fibers, glandular cells and other subdermal structures. Yet, in translucent specimens examined after
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minimal clearing, the presence of a relatively refractile, tubular struc- ture, about 25 /x in length is to be made out. This structure is in favor- able specimens quite discrete, and is to be interpreted as a vestigial or rudimentary buccal capsule; it sometimes appears to be protrusible and may function as a piercing organ or stylet. Its presence is de- scribed or illustrated in some of the other species which we attribute to this genus (e.g. Parlitoviosa zakii Nagaty), and upon its existence depends our views as to the taxonomie affinities of the genus.
Fig. 13. T etrapetalonema digitata. lateral views.
Posterior extremity of female; ventral and
The alimentary canal throughout its length is weakly developed. It is especially difficult to describe the oesophagus (which is only about 20^1 wide) when the specimens are completely cleared in lacto-phenol solution or glycerin. Faust's and McCoy's descriptions omit all reference to the alimentary canal. Chandler described the oesphagus of the species digitata as "very fine and slender, only about 11/x in diameter and not easily observed except where it is bent and runs at an angle to the long axis of the body. The junction with the intestine is not sharply demarcated, the intestine tapering, cone-like, to the junction with the oesophagus." Only by attempting to follow the course of the lightly cuticularized lining of the oesophagus to the point where it appears to end and where the coarser granulation of the intestinal cells commences does it appear possible to determine the
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approximate length of the oesophagus in our material. In the females this point is more difficult to detect than in male worms because the muscular dilatation of the vagina obscures the oesophago-intestinal
®
(^
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>9
Fig. 14. Tefrapetalonema digitata. Caudal region of male, ventral view, show- ing distribution of papillae.
junction. Consequently measurements of the oesophagus are no better than approximations which can be accorded no critical values.
The vulva is not salient, but the massive vaginal chamber, or ejector, allows its position to be easily located. The vagina runs a rectilinear course posteriad for 2 to 3 mm. and then divides into two uteri that are packed with microfilariae. As expressed from the uteri the fully developed larvae measure in watery solution from 170 to 188 /i
54
bulletin: museum of comparative zoology
in length and about 4fx in width. They are without a sheath and the posterior region tapers to a blunt end. Owing to the lack of suitable
Fig. 15. Tetrapetalonema digitata. Caudal region of male, lateral view, showing characteristic shape of the spicules.
blood smears we cannot describe the finer details of the microfilaria morphology.
The anus in the female has a rudimentary appearance indicating, perhaps, functional atrophy. The tip of the tail has a shallow cleft,
sandground: parasitic worms 55
and is ornamented by a pair of cuticular lappets, about 4/i in length, which originate on the sub ventral surface (fig. 13) .
In the preserved state the tail of the male is disposed in a loose two- looped spiral. The structure of its tip is essentially the same as de- scribed for the female. There are ver}^ narrow lateral expansions of the cuticle suggesting caudal alae, but these appear to be artefacts which are visible only when the tail is flattened by pressure from the cover- glass. The caudal papillae are very minute, sessile and subventrally situated. Usually they number three pairs in the peri-cloacal zone, one pair mid-caudal in position and a fifth pair immediately in front of the terminal lappets. As in many filariid species, the caudal papillae are subject to a degree of numerical irregularity and they are often asymmetrically disposed (fig. 14).
The spicules have specifically distinctive forms illustrated in figure 15. The " shaft," or proximal cylindrical portion, of the left spicule is as long or slightly longer than its more filamentous, leash-like, distal portion. The right spicule resembles a sickle in having a well defined "handle" and an arcuate "blade." In none of our specimens did we find the spicules exserted so as to permit the examination of their extremities. In this connection it may be of interest to note that the relatively long, filiform, left spicule in other species of the genus always appears to be extruded in fixed material.
There is no trace of a cuticularized accessory piece or gubernaculum.
The dimensions of T. digitata are given in the appended table.
Taxonomic Relations
As mentioned in our introduction, prior to the present re-allocation of D. (?) digitata Chandler, three species had been definitely assigned to Tetrapetalonema which, imputed to be closely allied to the genus Dipetaloncma, has been referred to the Setariinae.
A survey of the recent helminthological literature allows us to trans- fer an additional three species to Faust's genus: (1) T. nicollei (Mazza, 1929) and (2) T. tenuis (Mazza, 1929) both parasites of Cebus spp. from the northern " chaco" of Argentina; (3) T. zakii (Nagaty, 1935), a parasite of the Brazilian Leontocebus rosalia.
It is not improbable that other parasites of primates incompletely described as species of Filaria by early helminthologists such as Molin may also be members of Tetrapetalonema. Furthermore, it is possible that Tetrapetalonema is a synonym of Mansonella Faust, 1929, whose type and only species, M. ozzardi (Manson, 1897) parasitic in man in
56 bulletin: museum of comparative zoology
the Neotropical region, has been very insufficiently described on the basis of the female worms, which are very tenuous and have tails carrying minute lappets resembling the structures found in Tetrape- talonema spp.
In describing Parlitoviosa zakii, a species that appears to be very closely related to T. marmosetae, Nagaty (1935) referred his new genus to the Filariinae and compared it with such genera as Litomosa, Mac- donaldius, Ackertia, etc. In a more comprehensive review than we are at present able to undertake, several of these recently proposed genera, whose differentiation is rather indefinite, will have to be taken into account.
In this paper our consideration will be confined to those species of Tetrapetalonema that have been named above. In so far as several of these have been described in journals that are not widely accessible, it has been deemed desirable to present in tabular form that facilitates comparison, such measurements of the species as have been provided in the original descriptions.
Among many groups of parasitic nematodes, particularly those belonging to the Filariidae, differences up to 100% or even more in the gross size of individual worms, supposedly belonging to a single species, are not rare. Even the dimensions of the spicules and the proportionate length of such organs as the oesophagus are often subject to considerable variation. The taxonomic significance of divergencies such as these has been well recognized by some helminthologists, but their evalua- tion is difficult. Only by securing material from controlled experi- mentally induced infections does it appear possible to determine whether these variations are inherent in the constitution of individual worms without reference to any influence that the environment (involving such factors as unfavorable host species, etc.) may exert upon growth, or whether the differences are due to random samplings of worms of different ages, whose growth does not cease with the attain- ment of sexual maturity. For the critical student of helminthology, at least a notion of the extremes in the variation range is desirable before commensural data can be accepted as valid differential criteria for species that have not been distinguished from their congeners on a structural basis of constant nature.
As the accompanying table shows, although the dimensions in some species of Tetrapetalonema show appreciable variation, no indication of the size range is available for at least four of the seven supposedly distinct species. On the basis of present figures, three species (namely: T. digitata, T. marmosetae and T. parvum) appear to be separable on
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commensural data. However, some of these species may be dis- tinguished on criteria that are of greater taxonomic value than length. Thus in our experience, T. digitata may be recognized by the char- acteristic form of both spicules in the male, the right spicule, for example, being sickle-shaped and more easily seen than in T. mar- mosetae, T. atalensis and T. parva. On the other hand the buccal vestibule in T. atalensis is a well developed organ of characteristic shape (figure 16) which may be contrasted with the cylindrical, more or
Fig. 16. Tetrapetalonema atalensis McCoy. Anterior extremity in lateral view showing buccal capsule.
less diaphanous buccal capsule in the species parva, marmosetae and digitata. Furthermore, according to McCoy, the size of their respective microfilariae affords another means of distinguishing between certain of the species.
Without recourse to authentically identified specimens it is not possible to venture an opinion on the status of T. zakii, T. nicollei and T. tenuis. The descriptions of the two last mentioned species leave much to be desired. Their taxonomic integrity, and, hinging upon this, the nomenclatural status of other species in the genus, including the genotype, will remain in doubt until more complete descriptions are forthcoming.
sandground: parasitic worms 59
BIBLIOGRAPHY
Chandler, A. C.
1929. Some new genera and species of nematode worms, Filarioidea, from animals dying in the Calcutta zoological garden. Proc. U. S. Nat. Mus., 75, Art. 6, p. 1.
Faust, E. C.
1935. Filarial infection in the Marmosets, Leontocebus geoffroyi (Pucheran) and Saimiri orstedii orstedii (Reinhardt) in Panama. Trans. Roy. See. Trop. Med. & Hyg., 28, p. 627.
Mazza, S.
1929. Doblo parasitismo por filarias en monos Cebus del Norte. Quinta Reunion Soc. Argentina Patol. Regional del Norte, p. 1140.
McCoy, O. R.
1936. Filarial parasites of the monkeys of Panama. Amer. Journ. Trop. Med., 16, p. 363.
Nagaty, H. F.
1935. Parlilomosa zakii (Filariinae) a new genus and species and its Microfilaria from Leontocebus rosalia. Journ. Egypt. Med. Assn., 18, p. 483.
Sandground, J. H.
1933. Report on the nematode parasites collected by the Kelley-Roosevelt expedition to Indo-China, with descriptions of several new species. Ztschr. f. Parasitenkunde, 5, p. 542.
60 bulletin: museum of comparative zoology
3. Setaria hyracis Baylis, 1932, a synonym of S. loveridgei
Sandground, 1928
In 1928 I described Setaria loveridgei n.sp. on the basis of a few filariid nematodes that Mr. Arthur Loveridge had collected in Tan- ganyika from the hyrax, Procavia brucei prittivitzi at Dodoma, Tan- ganyika Territory. Four years later, Baylis (1932) described what appeared to be a second species of this genus, S. hyracis from a tree dwelling hyrax, Dendrohyrax sp, of the Belgian Congo (Sakuru), differentiating his species from S. loveridgei on such criteria as the smaller size, longer spicules and differences in the number and arrange- ment of the caudal papillae in the male.
In more recent years I have had occasion to identify specimens of Setaria taken from Procavia brucei frommei, P. scheffleri and Dendro- hyrax (?) arhorcus collected in various localities in Kenya Colony, Tanganyika and the Belgian Congo. As a result of these studies I have come to the conclusion that all of the material from the above named hosts comprised a single but highly variable species, presum- ably to be identified with S. hyracis. However, a re-examination of the type specimens of S. loveridgei has shown that a serious error was made in my original description of the spicules of this species.
The material upon which the description was based was extremely dark in color and even bleaching in hydrogen peroxide failed to render the two male worms as translucent as was desirable. In the male worm that was chosen as type, the right spicule was partially exserted. Re- examination has shown that what I originally took to be two spicules closely applied to each other along one side, is actually nothing more than the exceptionally broad proximal part of the right spicule, whose distal extremity was broken off; the right spicule, situated a con- siderable distance further forward was overlooked on account of the poor visibility. A study of the second male specimen shows the spicules to be of essentially the same form and size (right; 0.25 mm.; left; 0.59 mm.) as described by Baylis for »S. hyracis and as found in specimens from other species of hyrax. With these specimens, also, jS. loveridgei coincides in the general pattern of male caudal papillae and the appearance of the tooth-like projections in the circumoral ring, although it should be noted that the variation in the appearance of this structure is comparable with that described for S. hornbyi by Thwaite (1927).
As with many other members of the Filariidae, great disparity is to be found in the length of specimens of Setaria loveridgei from different
sandground: parasitic worms 61
species of hosts. In our first lot of material from Procavia brucei, the two males measured respectively 135 and 160 mm.; from P. scheffleri male worms vary in length from 100 mm. to 115 mm.; a single male taken from Dendrohyrax (?) arboreus is no longer than 65 mm. In the absence of any other taxonomically significant differences in these worms, it appears safest to regard them as intra-specific variants.
BIBLIOGRAPHY
Baylis, H. a.
1932. A new nematode parasite from a Hyrax. Ann. & Mag. Nat. Hist., Ser. 10, 9, p. 120.
Sandground, J. H.
1928. Some new cestode and nematode parasites from Tanganyika Terri- tory. Proc. Boston Soc. Nat. Hist., 39, p. 131.
Thwaite, J. W.
1927. The genus Setaria. Ann. Trop. Med. & Parasit., 21, p. 427.
Bulletin of the Museum of Comparative Zoology V *^^^_ '2 1939 AT HARVARD COLLEGE "^
Vol. LXXXV, No. 3
STUDIES OF DEEP-SEA ANGLER-FISHES (CERATIOIDEA)
By Talbot Howe Waterman Member of the Society of Fellows, Harvard University
CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
June, 1939
PUBLICATIONS OF THE
MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD COLLEGE
There have been published of the Bulletin, Vols. I to LXXXV, No. 1, 2 & 3; of the Memoirs, Vol. 1 to LIV, No. 1, 2 & 3, and Vol. LV
The Bulletin and Memoirs are devoted to the publication of original work by the Officers of the Museum, of investigations carried on by students and others in the different Laboratories of Natural History, and of work by specialists based upon the Museum Collections and Exploration.
These publications are issued in numbers at irregular intervals. Each number of the Bulletin and of the Memoirs is sold separately. A price list of the publications of the Museum will be sent on application to the Director of the Museum of Comparative Zoology, Cambridge, Massachusetts.
Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. LXXXV, No. 3
STUDIES OF DEEP-SEA ANGLER-FISHES (CERATIOIDEA)
By Talbot Howe Waterman Member of the Society of Fellows, Harvard University
CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
June, 1939
No. 3 — Studies on Deep-sea Angler-fishes (Ceratioidea)^
By Talbot Howe Waterman^ I. An historical survey of our present state of knowledge
In 1833 a strange and theretofore unknown fish was thrown up on the beach near Godthaab, Greenland, after a severe storm. It was remarkable for its bulky rounded body lacking ventral fins, black skin sparsely set with large spine-bearing plates, and especially for the "frontal tuft" arising from a longitudinal trough on the forehead and terminating in a bunch of filaments. Captain-lieutenant Carl Holboll procured this fish, unfortunately badly damaged by birds and decay, and sent it to Prof. Joh. C. Reinhardt at Copenhagen. The latter in 1837 described the specimen, which was fifty -eight centimeters in length, as adequately as possible under the circumstances and named it HimantoJophus groenlandicus. Because of the lack of proper material for study, Reinhardt was not explicit in placing his type in any of the known groups of fishes, but his comparison of this new form with Antennarius and Lophius, especially with reference to the "Pande- dusk," or fontal tuft, of Himantolophus, which he thought to be the same structure as, but more highly developed than the first dorsal ray on the head of Lophius, indicated that he believed his species to be related to those fish which comprise the Order Pediculati (Regan 1912). Himantolophus groenlandicus was the first deep-sea angler-fish to be discovered.
Before 1845 the same Captain Holboll who had collected this Himantolophus had obtained from the deep-sea off Greenland two more new "barbugede Tudsefiske" (called bare-bellied by the Danes because ventral fins are lacking in all the known ceratioid pediculates). One of these two was described in 1844 by Prof. H. Kroyer as Ceratias holbolli, while the other remained unnoticed, at least in the literature, until Prof. Chr. Liitken rescued it from dusty oblivion and described it as Oneirodes eschrichtii in 1871. Today these first specimens are con- sidered members of three distinct families of the Suborder Ceratioidea.
After these first ceratioids from high latitudes were discovered, occasionally others were taken, usually only a single specimen, some- times taken floating helplessly on the surface, as was Melanocetus
' Contribution Number 211. Woods Hole Oceanographic Institution.
2 1 am very much indebted to Dr. H. B. Bigelow, Director of the Woods Hole Oceanographic Institution, for many valuable suggestions made throughout the writing of this paper; it is a pleasure to thank him for his great assistance.
66 bulletin: museum of comparative zoology
johnsonii Giinther with another fish two or three times as long as itself coiled up in its stomach (Regan 1913). It was not, however, until the great oceanographic expeditions of the last fifty years or so that these peculiar fish were taken in any numbers. By far the most notable contributions to our knowledge of the Ceratioidea have come from the Danish oceanographic expeditions on the "Dana" under the leadership of .Johannes Schmidt. The 1920-22 Expedition took more than three times as many specimens as had probably been caught altogether up to that time and more than doubled the number of known species (Regan 1925a, 1926), and the 1929-30 Voyage-around-the-world again more than doubled the described ceratioids (Regan and Trewavas 1932) bringing the total, including those more recently described by Parr (1934), Schultz (1934), Fraser-Brunner (1935), and Belloc (1938, p. 303-6) to one hundred and sixty-four species.^
Since about eight hundred and sixty specimens are recorded in the literature, this would average five specimens per species. The distribu- tion of specimens has not, of course, been so fortunate, for two of the eleven families, twenty of the forty-one genera, and one hundred (or 61%) of the species are known from single specimens only. A mere handful of species are known from a sufficient number of specimens to allow any accurate ideas to be formed on their vertical and geographical distribution, changes in various parts and proportions in different stages of development, variability within the species, and many other such fundamental problems. There thus remains a tremendous amount of further collecting and comparative study of many more specimens before any adequate generalizations may be made concern- ing the biological relationships of these weird deep-sea fishes.
It was believed from an early date that the ceratioids were deep-sea fish, "degenerate" relatives of Lophius piscatorius L., living on the bottom with habits generally similar to those of this latter form. A species of Lophius common in the Mediterranean was well known to the ancients by such apt names as /Jdrpaxos aXtevs, Rana marina, Rana piscatrix, and the like. From the time of Aristotle vivid ac- counts were written of how this sly fish would use its frontal filament, or illicium, for attracting prey. The following passage (p. 67) from Oppian's "Halieutica," which was probably written in the reign of Marcus Aurelius (Emperor 161-180 A.D.), is an interesting example.
Most of the Renaissance ichthyologists, Rondelet, Belon, Salvianus and others, have passages describing similar behavior in Lophius. Ulyssi Aldrovandi in his "De Piscibus" (1612, p. 454-70) gives a non-
1 Oneirodes biMosiis Chapman (Proc. U. S. Nat. Mus.. Vol. 86, p. 538, 1939) brings the total up to one hundred and sixty-five.
waterman: deep-sea angler-fishes 67
critical but encyclopedic resume of the earlier accounts of the matter. With the beginning of the "modern" scientific era, soon after the time of these men, such descriptions of the fishing frog's feeding habits were thrown into disrepute as being but entertaining fishermen's tales.
Hid in the Slime the Toad of Form uncouth (That Fifli is all one vaft extended Mouth ) Her tender Body wraps, on Prey intent, And filent there concerts the great Event.
What fofter Skin, and flower Pace deny,
Wife Forefight and fuccefsful Frauds fupply
Within her Jaws a flefliy Fibre lies,
Whofe Whitenefs, grateful Scent, and Worm-IikeJ
Size
Attradl the Shoals, and charm their longing Eyes. She to allure oft fliakes the tempting Bait j They eager prefs, and hurry on their Fate. But as they near approach, with fubtle Art The wily Toad contracts th' inviting Part j Till giddy Numbers thus decoy'd flie draws Within the Circle of her widen'd Jaws.
The Fowler thus the featherd Race deceives, And ftrows beneath his Snare the rifled Sheaves. The bufy Flocks peck up the fcatter'd Seed, Nor midft their Joy the fatal Engine heed ; Till with loud Clap the tilted Cover falls. And the dole Pit the flutt'ring Prey enthralls.
(Translation of Diaper and Jones, Oxford 1722, lines 153ff.)
Much more recently, for example, Goode (1881, p. 469) considered that the dorsal tentacle of Mancalias uranoscopus (Murray) was more likely to be a sense organ for the detection of prey rather than an organ to attract it. This idea has had some confirmation from the work of Brauer (1908, S. 103), who found papillae which he considered to be
I
68 bulletin: museum of compakative zoology
gustatory, since they were well supplied with nerve fibers, on the bait or esca of the frontal filament of Gigantactis. None the less, there would appear to be a considerable amount of truth in the old stories about Rana piscatrix since recent observations of this fish in its natural habitat such as those cited by Bigelow and Welsh (1925, p. 528) and those of Chadwick (1929) and Wilson (1937) on aquarium specimens indicate that a Lophius lying concealed on the bottom in- deed often does lure other fish towards its waiting mouth by waving back and forth over its head the first dorsal spine with its terminal tag-like expansion.
This is undoubtedly true also of Antennarius, the frog-fish, another well-known shore water pediculate. Louis L. Mowbray, Esq., Director of the Bermuda Aquarium, has many times over a course of years observed the angling activities of aquarium specimens of this genus (personal communication 1938). When other small fish were placed in its tank, the Antennarius would slowly maneuver into a position near these other fish and begin to "angle." This consisted of moving about the rod-like first spine of the dorsal fin with its fleshy terminal expan- sion in a characteristic manner which was dift'erent in some cases for dift'erent kinds of prey. For example, when one kind of fish was in the tank with the Antennarius, it might slowly draw the first dorsal ray through the water in such a way that the "bait" moved back and forth in a sweeping arc, but when another kind of fish was present, it might vibrate the first dorsal ray and thus merely shake the "bait."^ Any fish which ventured near the Antennarius to investigate the moving "bait" was drawn very quickly into the enormous mouth cavity of the frogfish by a sudden gulp.
Histrio, the closely related Sargassum fish, which has taken up the specialized habitat of floating Sargassum weed, has also been observed many times under similar conditions, but interestingly enough, this fish has never been seen to use its first dorsal ray as a lure for prey but appears rather to depend entirely in obtaining its food on stealth and its inconspicuous appearance, which resembles remarkably that of the Sargassum to which it clings.
Samuel Garman (1899) in his description of the Pediculati from the "Albatross" collection of deep-sea fishes introduced two useful terms in referring to the frontal filaments of the fishes in this order. The whole first dorsal ray with its terminal elaboration he called the illicium. This was a Latin word meaning lure or inducement; it was
1 Wilson (1937, p. 488) similarly describes two comparable types of angling movements in Lophius.
waterman: deep-sea angler-fishes 69
not, however, at all frequently used by known Latin authors, and I have not been able to find it once used or referred to in any biological context earlier than Garman's, although a number of Latin authors used the related verb forms such as that seen in the quotation from Rondelet given below. According to Dr. Thomas Barbour (personal communication 1938), who was at the time a student in Garman's laboratory, Garman himself believed that he was coining a new biolog- ical term in illicium. Regan (1912), recognizing its usefulness, used the word again, and since that time it has been generally employed.
Garman called the terminal elaboration of the first dorsal ray the esca; this was a Latin word meaning bait, which had frequently been employed since classical times in reference to the use to which Lophius allegedly put its frontal filament. Cicero (ca. 45 B.C.) used esca in a description of the fishing frog's feeding habits in his "De Natura Deorum" (Lib. II, No. 125) ". . . ad quas quasi ad escam pisces quum accesserint . . .", and nearly all of the Renaissance natural history writers whom Needham has called the macro-iconographers of the sixteenth century, employed the same word similarly. Rondelet, for example, in his "De Piscibus Marinis" (1554, Lib. XII, Cap. XX) used it in the following passage concerning Lophius: "... ante (oculos) propendent appendices duae . . . quibus veluti esca pisces mira solertia allicit et capit quod non solum gravissimo Aristotelis testimonio, sed etiam piscatorum multorum experientia comprobatum est." It would seem obvious that Garman, who was a fine scholar of the old literature, had the term in question from such sources. Esca has not been used in recent ichthyological writings except by Garman, but I feel that the word would be a useful and convenient one to rein- troduce.
Thus when Reinhardt (1837) described the illicium of Himanto- lophus as being the same as, but more highly developed than the fishing filaments of Antennarius and Lophius, he certainly implied its use as a lure. Liitken (1871), however, seems to have been the first to state this directly, and he even suggested that the shape of the esca and its tentacles in Oneirodes might resemble the head of a nereid worm. The fact that some parts of the esca in at least certain of the ceratioids were luminous was first mentioned by Willemoes-Suhm (1876) in his letters from the "Challenger." This luminescence was thought to make the esca effective in the "eternally dark abysses" of the oceans, where it could not otherwise be seen.
Brauer (1908) in the anatomical section of his classic work on the "Valdivia" deep-sea fish made histological studies of the illicium and
70 bulletin: museum of comparative zoology
its terminal light organ in Gigantactis and Dolopichthys (which latter Brauer identified as Oneirodes). The light organ was described as a spherical, hollow, glandular structure located within the swelling of the esca and surrounded by a layer of reflecting cells and a heavy black pigment capsule. The lumen of the gland was filled by small granular bodies which Brauer considered to be secretory granules produced by the gland cells, some of which could be seen in his preparations break- ing down and apparently releasing from their cytoplasm similar struc- tures into the cavity of the organ. This lumen opened into a second, more distal cavity which in turn communicated with the exterior by a canal opening on the mid-dorsal line of the swelling of the esca. Be- cause of its general structure and especially because of the presence of the external opening, Brauer believed this organ to be a "Spitzdriise," that is a gland whose secretions were forcibly ejected. Never the less, he could not find any muscles in the esca which would permit such an active emptying of the gland's lumen. Neither was he able to find any nerve supplying the light organ although, as mentioned above, branches of an illicial nerve were distributed to the apparently sensory papillae of the esca. A fairly good blood supply, on the other hand, was found running to the luminous gland.
Similar terminal glandular organs may be present on other median fin rays of ceratioids such as those described by Brauer in the caruncles of the dorsal fin in Cryptosparas. As was noted by Regan (1926, p. 11) large specimens of this same genus may even have such structures present on the rays of the caudal fin. Luminescence has been observed by several workers to occur in the escal light organ, and Beebe (1934, p. 191-2) observed light coming from swellings on the tips of the first three dorsal rays in a ceratioid seen by him from the bathysphere; whether all such swellings on ceratioid fins are luminous or not is not known.
Dahlgren (1928) without more than merely stating that such was the case, described from histological examination the granules filling the cavity of the escal light organ of a specimen of Ceratias as lumi- nous bacteria. He presumed that these organisms were living sym- biotically within the esca, supplying light for the fish's bait in return for nutriment provided by a sort of culture medium maintained within the luminous gland by the activity of the secretory cells. A similar type of symbiosis had previously been described for the light organs of three or four other fishes, in some of which the results were quite convincing since careful bacteriological methods had been employed. (Dahlgren gave several references; see also Yasaki and Haneda 1936).
waterman: deep-sea angler-fishes 71
This same sort of interesting relationship has been found to occur in a number of luminous cephalopods. Bacteriological investigations of the light organs of a considerable number of different kinds of these animals (Kishitani 1932, Herfurth 1936) have shown that only in the case of luminous organs which open to the exterior by a canal or pore were symbiotic luminous bacteria present in large numbers; those light organs which were closed to the sea water did not contain the sym- bionts. The light organs of all the fish which were believed to cultivate luminous bacteria in their lumina were also open to the exterior. Dahlgren's conclusions concerning the bacterial origin of the light in Ceratias were thus in general agreement with such studies. However, in the case of another fish, the macrurid, Malacocephalus laevis, in which the light organ was open to the outside the luminescence was apparently not a bacterial luminescence since microscopic study demonstrated that the granules present in the material filling the lumen of the gland were not bacteria (Hickling 1925). Furthermore, the test for luciferin and luciferase,^ "substances" known to be in- volved in the luminescent reaction of a number of different organisms was positive. These materials have never been convincingly detected in bacteria so that it would appear highly unlikely that they were being extracted from symbiotic luminous bacteria in this case. It is clear then that at least two methods of producing light are employed by fishes, and it is possible that combinations of the two may be found in the same animal, but only after many more, careful investigations of the various types of luminous fishes and their light organs will it be possible to draw any satisfactory general conclusions concerning these matters.
Several references in the earlier literature to cases where living cera- tioids were examined in the dark, state that the light produced by the escal light organ was apparently under "voluntary" control; this meant at least that the luminescence was not continuous. In 1936 I had the opportunity to observe a living Dolopichthys which was taken by the research ship "Atlantis" in the Sargasso Sea. Still feebly moving this specimen was quickly removed from the net and placed in cold water. In the ship's dark-room, I was able to observe the luminescence of the escal light organ during the ten minutes or so that the fish remained alive. This remained dark until the fish was quite actively stimulated mechanically. When it did luminesce, the light was confined within the esca, which was transparent and more or less free from pigment
1 See E. N. Harvey. 1935. Luciferase, the enzyme concerned in luminescence of living organisms. Ergeb. Enzymforsch. Bd. IV, S. 365-79.
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distally, and no trace of luminous secretion was observed to be extruded from the external pore of the organ. The light was bluish green in color like that of the majority of luminous organisms and lasted for five or six seconds beginning dimly and rising to a peak in a second or so, maintaining the maximum intensity for several seconds, and then slowly fading out.
If the luminescence of ceratioids is a bacterial luminescence, the question of the control of the light is a very interesting one, for lumi- nous bacteria under normal conditions luminesce continuously, unlike all animals which produce their own light. A simple method of control of the light of these symbiotic bacteria would be through the amount of oxygen available to the bacteria, which luminesce only under aerobic conditions. It is bacteriologically conceivable that anaerobic condi- tions could be easily developed within the lumen of the light organ which would, when they became completely anaerobic, extinguish the light entirely. Sufficient oxygen for luminescence (the amount nec- essary is extremely small as was shown, for example, by the experi- ment of Beijerinck (1902) in which sufficient oxygen for momentary luminescence of the previously dark bacteria was obtained in a suspen- sion of clover leaves illuminated by the light of a single match) could easily be supplied by a blood system such a Brauer (1908) described in the ceratioids he studied. The present very imperfect state of our knowledge of the physiology of bioluminescence makes it unwise to do more than outline a few of the more salient points in this connection. Certainly the problem of the control of luminescence in animals is one of the least understood, yet biologically one of the most interesting of the many aspects of bioluminescence.
The nineteenth century students of ceratioids believed that these fish were benthonic, like Lophius. Filhol (1885, p. 81) even has an imaginative picture of Mdanocetus johnsonii Giinther in which several specimens are shown buried up to their eyes in the mud with their illicia waving over their heads. The fact that the mouth in this species opens upwards at a sharp angle made it a bit difficult to make these fish look plausible buried tail-downward in the ooze of the ocean bottom. Garman (1899) in his discussion of the genotype of Dolop- ichthys remarked on the improbability of such fish being active swimmers and stated his belief that their peculiar body form, fin reduc- tion, and smooth skin were adaptations for life in the soft muds of the sea floor. It was Brauer (1906) who first showed that the ceratioids were bathypelagic rather than benthonic forms. He based his demon- stration mainly on the analysis of vertical closing net hauls made by
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the "Valdivia" and which showed at what depths in the water the specimens were fished; he furthermore pointed out that several cera- tioids which had been taken previously had been found with scopeline and other non-benthonic fish in their stomachs. It is generally believed today that, with the exception of the young, which have often been taken in shallower water, the deep-sea angler-fishes live bathypelagi- cally largely at depths ranging from 500-5,000 meters (Regan and Trewavas 1932, p. 12; Beebe 1937, p. 207).
The occurrence of dwarf males living attached to the relatively gigantic females which has been described in a number of deep-sea angler-fish is such a striking and bizarre phenomenon that it has be- come a part of everyday scientific knowledge. Nevertheless, no thoroughly satisfactory analysis of the few available facts concerning this interesting sexual relationship of the ceratioids has as yet been published. Regan and Trewavas (1932) give a good descriptive section on sexual dimorphism, however.
Saemundsson (1922) was the first to discover the attached males. He noticed two small fish grown fast to the belly of a large female Cer alias holbolli; he did not recognize them as dwarf males but thought that they might possibly be young of the same species since they showed certain well marked similarities to the larger fish. Three years later Regan (1925b) from a dissection of another specimen of C. holbolli to which a small fish was attached showed that this latter was a male with a fairly well developed testis and that the larger fish was a female.
Up to the present time fourteen specimens of these dwarf males have been discovered attached to twelve (out of the approximately five hundred and fifty known) female ceratioids. Most of these females had only one male grown onto them, but Saemundsson's specimen as mentioned above, had two, and one female of Edriolychnus schmidti (Regan and Trewavas 1932, plate IX, fig 2) had three of them. The males have been found attached head-on to the females by outgrowths from either the lower jaws or from both the upper and lower jaws to a papilla raised up on the surface of the body of the females. The place of attachment was apparently random since males have been found fastened to the belly, head, and even the preopercular spine of the females.
The female angler-fish so far found with attached males belong to four distinct families of the Sub-order Ceratioidea. Regan (1925, p. 397) at first believed that attached males were characteristic of the whole sub-order; this point of \'iew was also favored by Parr. The latter worker, however, (1930) after a study of the anatomy of the
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then rather anomalous family of eeratioids, the Aceratiidae, was led to conclude that these small fish, lacking an illicium externally at least (except for Anomalophryne Beebe, 1929) and generally possessing pincer-like rostral denticles, large nostrils and olfactory organs, were free-living stages of male eeratioids theretofore recognized only in the attached forms. A careful study of the viscera of one of these free- living non-illiciate fishes, RhyuchoceraHas longipinnis, was made by Parr (1930). This specimen was indeed a male and had a testis ap- parently as large relatively as any found in the attached males which have been investigated in this respect. Determination of the sex of as many of the aceratiids as possible has led to the discovery that they were all males; furthermore, similar study has shown all the illiciate "typical" eeratioids to be females. From anatomical evidence it has been possible in most cases to assign the free-living males to families previously known only from females. Free-living males then have been found to occur in five of the eleven families of the Ceratioidea.
Parr (1930, 1932) suggested that the free-swimming males were doubtless merely at an earlier stage in their life history than were the attached males and would consequently become attached at some later time. On the other hand, Regan and Trewavas (1932, p. 20) thought it probable in view of Parr's discovery of the free-living males, that attached males occur only in the four families in which they had been found up to that time and that all the other families of eera- tioids had free-living males. This point of view was not sustained by any direct evidence presented but seemed rather to rest on Regan's (1925b, p. 396) assumption which he had maintained from the begin- ning that the attached males became fastened to the females at an early stage, probably shortly after the larval stage, certainly before the gonads have matured.
Evidence for a long period of attachment at least was found (Regan and Trewavas 1932, p. 42) in the skull of an attached male Ceratias holholli in which certain osteological peculiarities were believed to have resulted from the male's having been much smaller at the time of attachment. However, if Regan's original assumption corresponds to the facts then no free-living males with as well developed testes as Rhynchoceratias could be found in a family which had parasitic males. Yet free-living males have been found in the Linophrynidac. Thus the situation becomes rather confusing in the paragraphs concerning the male linophrynids (Regan and Trewavas 1932, p. 15fF) since both free- living and attached males are described in this family. The free- swimming males of the type of Aceratias were shown to be very similar
waterman: deep-sea angler-fishes /o
to the attached males of the Genus Borophryne. In spite of the fact that the free-living male Linophrynidae listed by Regan and Trewavas (1932) average over twenty-five percent longer in standard length than the attached males discussed, these authors felt obliged to main- tain without qualification the assumption of an early time of attach- ment and the double hypothesis that free-living males occur in certain families of ceratioids and attached ones occur only in the remaining families. These views would seem to be rather troublesome to main- tain especially since Parr's (1934, p. 56fF) discovery of a free swimming male, Borophryne masculina, belonging to a genus in which two females with attached males are already known. In this case, too, the free- living male is somewhat larger than the attached ones of the same genus.
It would thus seem clear that the simplest reasonable explanation of the facts would be that free-living and attached males might well be found to occur in all of the deep-sea angler-fish which have developed a marked sexual dimorphism. Furthermore, as Parr has pointed out (1930, p. 134), the males do not generally become attached at an early stage of their development since, for example, amongst the Lino- phrynidae mentioned above there are one hundred and thirty free- living males recorded in the literature and only eight attached males. The time of attachment and consequently the period during which this union of male and female lasts would appear from the evidence re- viewed to vary with the individual circumstances.
It has been generally assumed that the development of males which were first free-swimming fairly active fishes and then sessile on the females was an adaptation to the fact that the female ceratioids were solitary sluggish animals whose chances of mating in the dark abysses of the oceans would be rather negligible if the males of the group were of similar habit. This adaptational point of view would go on to sug- gest that if a male and a female of the same species were so fortunate as to find one another, some means of their continued association until their reproductive functions had been discharged would be of great advantage. Thus it may be that certain of the individual males find and attach on to their mates soon after the post-larval period, whereas others swim about seeking mates until they have become mature, and doubtless still other males never do find a female of their species. Also it may be in the more primitive families of the sub-order, in which no attached males have yet been found, that instead of actually grow- ing fast to the females, as was the case of the attached males already known, the males merely nip on to the females for a relatively short
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period. The fact that only two percent of all the known female cera- tioids and only seven percent of the females of just the families in which attached males have already described have males attached to them may be indicative of a relatively brief period of union between the two sexes. On the other hand, it may rather indicate, in conjunc- tion with the very small size of the great majority of the angler-fish which have yet been captured, that only a relatively few mature fish have been taken so far.
Another point of interest concerning the male ceratioids is that although fifty-seven percent as many free-living male specimens as females have been found, there are only twenty percent as many species of males as there are female species. Thus there are not nearly enough species of males to pair them up with the females. If all the specimens which are assumed to be female are, it would seem likely then that either differences which were either individual or ontogenetic have been used in defining many of the female species, or else the ability to differentiate between males of different species has not been sufficiently developed to distinguish the rather obscure details which separate them.
The fact that both the Melanocciidae and the HimantoJophidae, the most primitive familes of the Ccratioidea, have two genera of males to one of females has been pointed out by Regan and Trewavas (1932, p. 21) to be in striking contrast with the rest of the sub-order. The fish in one of the male genera in both cases, Centrocetus and Lipactis, were more like females in shape than any other known free-living males; teeth in the jaws, which were generally absent in male cera- tioids, were present, and the rostral denticles were not well developed. The more typically ceratioid appearance and the very small size of these peculiar males would suggest the possibility that they were merely juvenile specimens. A minute male of the Family Dicrra- tiidac, Caranadis pumilus, would presumably belong in the same category.
The physiological significance of the attachment of the males to the females in the deep-sea angler-fish is a point of great interest. Regan from his first paper on the subject has maintained that the attached males were nutritional parasites on the females. Sections cut through the region of attachment of the two fish were believed to demonstrate a complete blending of the vascular systems in that region indicating that they were continuous from one fish to the other (Regan 1926, p. 12). F'urthermore, the mouth of the males, because of the nature of the attachment to the female, was no longer available for active feed-
waterman: deep-sea angler-fishes 77
ing, the stomach and gut were remarkably degenerate, and the union between the two was beheved to last for a long time. All of these con- ditions would seem to indicate that the males must depend on some unusual source of nourishment. Further descriptions and figures of the region of fusion between the male and female, as it appeared in serial sections, were given by Regan and Trewavas (1932, p. 21); sinuses of the region were traced out as connecting the blood vessels of the pair through a capillary network on each side, thus proving the direct continuity of the blood streams that had been strongly sug- gested by previous studies. Parr (1930, 1932), however, expressed the opinion that the nutritional aspects of the continuity of the circulatory systems might not be as important as previously believed. He was led to this point of view, among other things, by the relatively enormous size of the liver in the specimen of free-living male whose anatomy he had investigated. Thus before the time of its attachment to the female, the male might store up a large quantity of food in its liver, which would be an efficient storage space; this food could then be used by the male during the period of his union with the female. Evidence in support of this suggestion may be obtained from a number of figures in Regan and Trewavas (1932, pp. 16, 17, 20, 22) of the visceral anat- omy of male ceratioids. These figures show that the free-living males had relatively large livers whereas in the attached males which have been examined the livers were small. It is interesting to note in this connection that the attached male of Ceratias holholli which Regan and Trewavas believed from osteological evidence to have been fast- ened to the female for a long period had apparently an extremely minute liver since this organ was neither shown in the figure of the anatomy of this fish nor mentioned in the text (Regan 192ob, fig. 4 and p. 392). Perhaps in cases where the attachment of the male lasts over a longer period than could be supplied with food stored in the liver, the nutritional possibilities of a confluence or even placenta-like arrangement of the blood streams may come to be realized.
A further structural arrangement pertinent to these matters has been known from the time of Saemundsson's (1922) discovery of the first attached male. Namely, the fact that in every case although the mouth of the attached fish becomes more or less occluded from the nature of the attachment, lateral openings into the mouth cavity are retained. These are believed to permit the circulation of a respiratory current. Regan (1926, p. 14) stated that since the attached males have a well developed heart and gills, they do not depend on the blood of the females for oxygen but only for nutritive materials. As Parr
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(personal communication 1938) has pointed out, if arrangements for a respiratory current and normally developed respiratory and circula- tory systems are maintained by the attached males, they could not be receiving sufficient oxygenated blood from the females for their own, probably meager, needs. Either then the male receives only venous blood from the female or else not much blood at all. Two facts would ap'pear to cast some light on the probability of the former alternative : First, Regan (1926, p. 14) states that both arteries and veins run into the capillary network of the tissues connecting the males and the females. Second, if the males were carrying the added excretory burden of a considerable venous circulation from the females, unusually large kidneys might be expected in the attached males; these organs have not been found to be of greater size, however, than those found in the free-living males. If these arguments carry any weight, then we are left with the alternative that there is not much blood interchange between the males and the females.
One idea that has been suggested by both Regan and Parr is the possibility that the continuity of the blood streams would permit an hormonal interchange between the male and the female which might affect the differentiation of the gonads, and synchronize their maturation and the discharge of their sexual products. However, from the general results of vertebrate endocrinology, especially as related to parabiosis, it would appear unlikely that the attachment of the males, provided they were sexually differentiated (and there is no mention in the literature of any attached "male" in which the gonad could not be easily identified as testis) and normal as to their endocrine functions, was of any particular significance (Hisaw, personal com- munication 1938). It must be admitted, never the less, that in animals which have become morphologically as highly specialized as the ceratioids it would be difficult to predict with any degree of assurance what special physiological functions they may have developed in the course of their evolution.
The foregoing account has attempted to outline and assay the facts which are known concerning the deep-sea angler-fish. In the analysis of even the most recent knowledge of these fish the discussion has been filled with probabilities, suggestions, ideas, possibilities, and the like, and nearly every problem approached was found to rest in a highly unsettled and hypothetical state. This at the present time is true of our biological knowledge of all bathypelagic organisms. The great difficulty and expense of obtaining material and the well nigh in- superable barriers to experimental work in this field have been the
waterman: deep-sea angler-fishes 79
principal deterrents to great advances in these directions. It is to be hoped, however, that in the future with the further cooperation of the great oceanographic institutions and their research vessels that many of these intriguing and significant problems may be satisfactorily solved and fitted into the general body of physical and biological knowledge which constitutes the science of oceanography.
BIBLIOGRAPHY
Beebe, W.
1929. Ilaplophryne hudsonius. A new species; description and osteology. Zoologica, Vol. 12, p. 21-36.
1934. Three new deep-sea fish seen from the bathysphere. Bull. N. Y. Zool. Soc, Vol. 37, No. 6, p. 190-3.
1937. Preliminary list of Bermuda deep-sea fish. Zoologica, Vol. 22, p. 197-208.
Beijerinck, M. W.
1902. Photobacteria as a reactive in the investigation of the chlorophyll function. Proc. Roy. Acad. Amsterdam, Vol. 4, p. 45-9
Belloc, G.
1938. Liste des poissons pelagiques et bathypelagiques captures au cours de la cinquieme croisiereavec diagnoses preliminaires de deux especes nouvelles. Rev. Trav. Office des Peches, T. XI, fasc. 3, p. 281-313.
BiGELOw, H. B. and W. W. Welsh
1925. Fishes of the Gulf of Maine. Bull. U. S. Bur. Fisheries, Vol. 40 (1924), Pt. 1, p. 1-567.
Brauer, a.
1906, 1908. Die Tiefsee-Fische. Lief. I, Systematischer Teil, S. 1-420. Lief. II, Anatomischer Teil, S. 1-266. Wiss. Ergeb. Deut. Tiefsee-Exped. . . . "Valdivia," Bd. 15.
Chadwick, H. C.
1929. Feeding Habits of the Angler-fish, Lophius piscatorius. Nature, Vol. 124, p. 337.
Dahlgren, U. •
1928. The bacterial light organ of Ceratias. Science, N. S., Vol. 68, p. 65-6.
FlLHOL, H.
1885. La vie au fond des mers. Paris.
Fraser-Brunner, a.
1935. New or rare fishes from the Irish Atlantic slope. Proc. Roy. Irish Acad., Vol. 42B, p. 319-26.
80 bulletin: museum of comparative zoology
Garman, S.
1899. Reports on an exploration ... by the "Albatross" during 1891. XXVI, The Fishes. Memoirs Mus. Comp. Zool., Vol. 24, p. 1-431.
GooDE, G. B.
1881. Fishes from the deep water on the south coast of New England obtained by the U. S. Fish Commission in the summer of 1880. Proc. U. S. Nat. Mus., Vol. 3 (1880), p. 467-86.
Herfurth, a. H.
1936. Beitrjige zur Kenntnis der Bakteriensymbiose der Cephalopoden. Zeit. Morph. Okol. Tiere, Bd. 31, S. 561-607.
HiCKLING, C. F.
1925. A new type of luminescence in fishes. J. Mar. Biol. Assoc, N. S., Vol. 13, p. 914-37.
KiSHITANI, T.
1932. Studien iiber Leuchtsymbiose von japanischen Sepien. Fol. Anat. Jap., Bd. 10, S. 315-418.
Kroyer, H.
1844. Meddeler Forogelser til Kundskab om de gronlandske Fiskearter. Overs. K. D. Vidensk. Selsk. Forh. 1844, No. 8, S. 138-41. (Probably published before the following; Ceratias holbolU here nomen nudum.)
1844. Ceratias Ilolhdlli Kr. Naturh. Tidsskr. Bd. 1, Ser. 2, S. 639-49.
Lutken, C.
1871. Oneirodes Eschrichtii Ltk. en ny gronlansk Tudsefisk. Overs. K. D. Vidensk. Selsk. Forh. 1871, S. 56-73. (An Enghsh translation of this paper appeared in Ann. Mag. Nat. Hist., Ser. 4, Vol. 9, p. 329-44, 1872).
Parr, A. E.
1930. On the probable identity, life-history, and anatomy of free-living and attached males of the ceratioid fishes. Copeia, 1930, p. 129-35.
1932. On a deef)-sea devilfish from New England waters and the peculiar life and looks of its kind. Bull. Boston Soc. Nat. Hist., No. 63, p. 3-16.
1934. Report on experimental use of a triangular trawl for bathypelagic collect- ing. Bull. Bingham Oceanogr. Coll., Vol. 4, Art. 6, p. 1-59.
Regan, C. T.
1912. Classification of the teleostean fishes of the Order Pediculaii. Ann. Mag. Nat. Hist., Ser. 8, Vol. 9, p. 277-89.
1913. A deef)-sea angler-fish, Melanocetus johnsonii. Proc. Zool. Soc, London, 1913, p. 1096-7.
1925a. New ceratioid fishes from the North Atlantic, the Caribbean Sea, and the Gulf of Panama, collected by the "Dana." Ann. Mag. Nat. Hist., Ser. 9, Vol. 15, p. 561-7.
waterman: deep-sea angler-fishes 81
1925b. Dwarfed males parasitic on the females in oceanic angler-fishes {Pedi- culati Ceratioidea) . Proc. Roy. Soc. Lond., Vol. B97, p. 386-400.
1926. The pediculate fishes of the Suborder Ceratioidea. Danish "Dana" E.xped. 1920-22, Rep. No. 2, p. 1-45.
Regan, C. T. and E. Trewav.\s
1932. Deep-sea angler-fishes (Ceratioidea). Carlsberg Foundation "Dana" Report No. 2, 1932, p. 1-113.
Reinhardt, J. C.
1837. Ichthyologiske Bidrag til den Gronlandske Fauna. K. Danske Vidensk. Selsk. Nat. og Math. Afh., 4te Raekke, Bd. 7, S. 83-196.
Saemundsson, B.
1922. Zoologiske Meddelelser fra Island, XIV. 11 Fiske, ny for Island, og supplorende Oplysninger om andre, tidligere kendte. Vidensk. Medd. Kobenhaven, Bd. 74, S. 159-201.
SCHULTZ, L. P.
1934. A new ceratiid fish from the Gulf of Alaska. Copeia, 1934, p. 66-8.
Willemoes-Suhm, R. aon
1876. Challenger Briefe an C. Th. E. von Siebold. Zeit. Wiss. Zool., Bd. 26, S. Ixxxi.
Wilson, D. P.
1937. The habits of the Angler-fish, Lophius piscatorius L., in the Plymouth Aquarium. J. Mar. Biol. Assoc, N. S., Vol. 21, p. 477-96.
Yasaki, Y. und Haneda
1936. Uber einen neuen Typus von Leuchtorgan im Fische. Proc. Imp. Acad. Tokyo, Vol. 12, p. 5.5-7.
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11. Three new species
In the course of examining hauls made from the research ship "Atlantis" during the summers of 1936 and 1937 a few small Ccrati- oidea were found. Three of these are apparently new species belonging to, 3 distinct families of this suborder: Gigantactinidae, Linophrynidae, and Oneirodidae.
GiGANTACTis LONGiciRRA, spec. nov. (Figs. 1 and 2)
1 specimen: holotype, M C. Z. No. 35,065.
Total length about 62 mm., standard length 39 mm. Illicium ter- minal on snout, length from articulation with basal bone to distal end of the bulb 15 mm. (39% of S. L.). Length of lower jaw 7 mm. (18% S. L.); eye diameter O.S mm. (2%o S. L.).
Fin rays. D.8, A. 6, C.8, P. 18. Branchiostegal rays 6 (2+4) ; pectoral radials (pterygials) 5 (4+1).
Dentition. All teeth curved and depressable inward or backward. Lower jaw: 13-14 teeth on each side, of different sizes, ranged in 2 rows one within the other. Upper jaw: 8 in a single row on each premaxil- lary; maxillary toothless. Lower pharyngeals toothless; upper pharyn- geals: 2nd and 3rd pharyngobranchials each have 3-4 large teeth. Palatines and vomer toothless.
The esca"^ (Fig- 2) consists of a somewhat pear-shaped swelling, rounded and blunt distally, and bearing at its proximal end 20 slender filaments which are 60-300% as long as the swelling itself. Each of these filaments terminates distally with a just perceptible bulbous expansion. Within the pear-shaped swelling is a heavily pigmented spherical core (the light organ) with a hemispherical prominence of about half its own diameter projecting from it postero-dorsally. The core is borne on the larger end of a conical expansion of the inner stalk which runs up the center of the illicial stem.
The nostrils are tubular; lateral line organs on projecting tags or flaps of pigmented skin are present on the head, body, and also the rays of the caudal fin. There are no channels or grooves connecting these organs with one another. The skin is closely set with small sub- circular plates each bearing a short sharp spine.
No pelvic bones were found in a dissection of one side of the speci- men.
' For definition and origin of this term and "illicium" see preceding section, p. 68-9.
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83
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Color. Chocolate brown (after fixation in formalin, followed by alcohol.)
Type locality. "Atlantis" Station 2894, July 20th, 1937, continental slope about 300 miles east of C ape May, New Jersey (39°06'N., 70° 16'W.), closing net at 1000 m., 4 A.M., depth to bottom 2860 m.
Discussion
Clearly the slender, elongate body of this ceratioid, the attenuate frontal region of its skull, and the long illicium located terminally on the snout demonstrate that it belongs to the Gigantactinidae Bou- lenger, 1904, and more specifically to the genus Gigantactis Brauer, 1902. Of the 8 individual specimens of the genus, which have been taken previously, 7 are types of as many different species (Brauer, 1902; Regan, 1925, 1926; Regan and Trewavas, 1932; and Fraser- Brunner, 1935) from all of which the present specimen is notably distinct. Four characters clearly set it apart from any other known form. These are:
1 . The number of dorsal rays is greater by 2 than that given for any of the other Gigantactinidae ; all other members of the present genus have 6 and Rhynchactis leptonema 3 (Regan, 1925)^
2. In the pectoral appendage one more pectoral radial bone is present than has been described for any of the other Pediculati, which primitively have only 3 of these bones (Gregory and Conrad, 1936, p. 202). Regan and Trewavas (1932) have found 4 of these elongate basalia in both G. se.rfilis and Rhynchactis leptonema and suggested (p. 25) that the presence of 4 of them is a diagnostic feature for the Gigantactin idae.
3. The length of the illicium from its proximal end, where it articu- lates with the modified pterygiophore or "basal bone," to the distal end of the bulb of the esca is less than 40% (38%) of the standard length. In previously known species, including smaller as well as larger specimens, this measure was always 60% or more of the standard length, varying from 60%; in G. exodon (where it might be a juvenile character since the type is only 25 mm. in standard length) to 400% in G. macronema.
4. The esca is notably different from that described for any other ceratioid. Except for exodon, all other gigantactids have terminal filaments present on the esca whereas in the present fish there is no
■ Only 5 dorsal rays are shown in the figure of G. vanhoeffeni although 6 is the number given in the description (Brauer, 1906).
waterman: deep-sea angler-fishes 85
trace of such. In exodon, however, the swelHng of the esca tapers dis- tally; this is definitely not the case with the new species. Yet in the only other species of Gigantactis, ovifcr and fiUbulhosus, in which the bulb does not taper distally there are two terminal filaments present in the esca and no proximal filaments at all. The proximal filaments of the new species' esca are more numerous and longer proportionately than any such filaments found in other specimens of Gigantactis.
A further point of interest in G. longicirra concerns the lateral line organs. In other members of the family (Regan and Trewavas, 1932, p. 24) these occur on "stalks" rather than on "tags" as found in longicirra. If this difl^erence does not depend on confusion of ter- minology it may well be a fifth clear distinction of the present species. Brauer (1906) figures G. vanhoeffeni with lateral line organs on the caudal rays much as in longicirra but does not mention this unusual phenomenon in the text.
Although the standard length of the present specimen is only 40% of that of the largest specimen of Gigantactis yet taken (the type of G. macronema Regan, 1925), one half of the types of previously known gigantactinids are smaller. Thus it would appear that the characters used to distinguish G. longicirra are no more likely to be juvenile than are those for half the members of the genus. Furthermore, it would not seem likely that the structural characteristics listed above (at least the first two) had been derived by growth or simplification as a stage in the life history of some previously known form. However, in the ab- sence of any direct evidence concerning the changes which take place in Gigantactis in the course of its ontogeny, it is rather difficult to make any predictions concerning such aberrant forms. ^
Linophryne algibarbata, spec. nov. (Figs. 3 and 4)
1 specimen; holotype, M. C. Z. No. 35,066.
Total length 40 mm., standard length 28 mm. lUicium suprarostral, length from articulation with basal bone to distal end of bulb 7 mm. (25% of S. L.), basal bone not exserted in present condition of speci- men.- Bulb of esca 50% of length of illicial stem. Length of lower
' But see Fraser-Brunner's (1935) suggestion that the presence of teeth on the lower jaw well down on the sides of the chin and away from the bite of the jaws was a juvenile character in Gigantactis.
2 Since the basal bone (modified pterygiophore) of the iUicium is freely naoveable and supplied among others with extensor and retractor muscles, the amount of its exsertion in any specimen depends on the condition of these muscles on fixation. Clearly, however, certain forms such as Mancalias and Lasiognathus must always have a considerable portion of the basal bone exserted whereas others as Edriolychnus and Himantolophus do not.
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jaw to articular spine 9 mm. (31% S. L.). Barbel length about 24 mm. (86% S. L.).
Sphenotic and preopercular spines large and sharp, free of skin except at their bases, smaller frontal and articular spines present.
Fig. 4. Linophryne algibarbaia, spec. nov.
Dentition. Teeth depressable inwards. Lower jaw: 7-8 irregularly arranged teeth including 1 very large (15% S. L.) and 2 smaller fangs on each side. Upper jaw: 7-8 irregularly arranged teeth including 1 large fang on each premaxillary; maxillary toothless. Large fangs saber-like, sheathed proximally in pigmented epithelium. Lower
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pharyngeals toothless, 3 teeth each on second and third upper pharyn- geals (pharyngobranchials). Palatines toothless; 2 teeth on vomer.
Fin rays. D.3, A. 2, P. 16. Brachiostegal rays 4 (1+3); pectoral radials (pterygials) 3 (2+1). Pigmented skin runs out over all fin rays.
Barbel arises from short stalk which divides immediately into four main branches of approximately equal length and diameter; each of these gives rise to irregularly arranged, much shorter secondary and tertiary branches, which in most cases have knob-like swellings along their lengths and terminally.
Esca consists of a subspherical bulb bearing around its anterodorsal surface a roughly horse-shore shaped ridge which consists of a median and two lateral lobes. In the median line, just posterior to this ridge is a low silvery papilla behind which is the external pore of the luminous organ of the esca. A slight longitudinal ridge of short extent is present immediately posterior to this. A black pigment cup which is visible through the transparent outer layers of the bulb surrounds about half of an inner sphere (the light organ), the exposed, distal part of which appears silvery. The rim of this pigmented cup has two distinct notches in it, one anterior, which is visible in Fig. 4, and one posterior.
The nostrils are large and tubular; lateral line organs are present on the head and body; these are situated on tags of pigmented skin, a few of which are notably larger than the rest. No channels or grooves con- nect these organs.
No pelvic bones were found in a dissection of one side of the speci- men.
The skin is smooth and (after fixation in formalin followed by alco- hol) chocolate brown in color.
Type locality. 39°06'N., 70°16'W., July 20th, 1937; closing net, 400 m., 5 A.M., depth to bottom 2860 m.
Discussion
In recent papers Regan and Trewavas (1932) and especially Parr (1934) have organized and extended our knowledge of the genus Linophryne. Twenty-five specimens belonging to this genus have been reported in the literature. For the twenty-one of these which are adequately known thirteen species and five subspecies have been de- fined. Thus nine species and two subspecies are known only from single specimens. In no case has more than one fish of a species in this genus been taken in a single catch.
waterman: deep-sea axgler-fishes 89
The characteristics which distinguish L. algibarbata most clearly from any of these previously known species are as follows :
1. The absence of either terminal or lateral filaments or of digitiform appendages on the esca. These occur in all other known species in some form or combination.
2. The general pattern and the method of branching of the hyoid barbel. While this appendage is similar in a general way to that of L. arborifera and of L. macrodon, it differs distinctly from these not only in the fact that the four main branches are of approximately equal length and diameter but also in the details of the secondary and tertiary ramifications.
3. Presence of only 4 branchiostegal rays instead of the 5 which Regan and Trewavas (1932) had concluded from their dissections of L. racemifera, Halpophryne sp., and Aceratias indicus were char- acteristic of the Family Linophrynidae. However, the present observa- tion shows that more specimens will have to be investigated before the significance of this point for diagnostic purposes will be apparent.
Since the present specimen differs from forms previously described in the fundamental pattern of the barbel and of the terminal append- ages of the esca, it is clear that a new species is necessary here rather than a subspecies, such as those set up in Linophryne arborifera and L. coronata by Parr (1934) on the basis of minor variations in the details of these structures.
DoLOPiCHTHYs ALBiFiLOSA, spcc. nov. (Figs. 5 and 6).
1 specimen; holotype, M. C. Z. No. 35,067.
Total length 45 mm., standard length 33 mm. Illicium suprafrontal, length from articulation with basal bone to distal end of escal bulb 14 mm. (41% S. L. or 2.4 times as long as the exserted part of the basal bone in the present condition of the specimen')- Bulb of esca about 20% of length of illicium from basal bone articulation. Length of lower jaw 14 mm. (41% S. L.); lower jaw with symphysial spine. Large sharp sphenotic spines present; also 2 articular spines on each side, less than half the size of the sphenotics.
Fi?i rays. D.5, A.5, P.17. Branchiostegal rays 6 (2+4); pectoral radials (pterygials) 3 (1+2).
Dentition. Teeth slender and acicular, depressable inwards. Lower jaw: 24 teeth on each side arranged in three series of different sizes. Upper jaw: 24 in a single series on each premaxillary (only 7-8 of these evident without partial dissection); maxillary toothless. Up-
' See footnote 2, p. 85.
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per and lower pharyngeals and palatines toothless. Two teeth on each side of vomer.
The esca' (Fig. 6) consists of a laterally compressed terminal expan- sion of the illicium enclosing within it an ovoid bulb. On the median anterior aspect of the latter is a low papilla projecting anterodorsally. Just posterior to this and running laterally are two slightly raised areas; between these and in the median line is a ridge which runs posteriorly and bifurcates over the posterior portion of the bulb. Each limb of this bifurcation gives rise to a stout, rapidly tapering filament one-fourth as long as the illicium from basal bone articulation to distal end of bulb. A loose transparent layer of tissue covers the distal part
Fig. 6. Dorsal view of esca of Dolopichthys albifilosa, spec. nov.
of the bulb and terminates posteriorly in a little upstanding ruffle on the antero-median surface of each of the escal filaments. Posterior to the latter and in the median sagittal plane the escal bulb bears a large vane-like, laterally compressed appendage three-tenths as long as the illicium. Distally on the ventral margin of this appendage there is a subspherical heavily pigmented body with a ventrally directed white spot (light organ?). The remaining pigmentation of the whole struc- ture is as follows: the inner bulb of the esca, the proximal part of the latter where it expands from the illicial stem, and the median ridge running posteriorly from the papilla, are all heavily pigmented. The two raised areas just behind and lateral to the papilla, the ventral part of the compressed appendage and the anteromedian aspect of the escal filaments are lightly pigmented by a few scattered granules. The remaining parts of the esca including the rest of the two filaments are whitish.
The nostrils are short and tubular with two openings distally ; lateral
• The present description of the esca is made with the orientation of this organ in the living animal in mind. "Posterior" and "anterior" are reversed in relation to Regan and Trewavaa' (1932) use of these terms.
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line organs on tags or flaps of pigmented skin with terminal white spots are present on the head and body. An unusual aggregation of these organs occurs caudally between the opposed soft dorsal and anal fins.
A pelvic bone was found in a dissection of one side of the specimen.
The skin is smooth and jet black.
Type'locaUhj. "Atlantis" Station 2667 (35°40'N., 69°36'W.), depth of haul unknown, September 1936.
Discussion
It is clear that the present specimen is closely related to Dolo- pichthys mgrifiUs Regan and Trewavas (1932, p. 67). There are, how- ever, a number of distinct differences between them :
1. The escal filaments of D. alhifilosa are almost entirely free of pigment, whereas the proximal four-fifths of the same appendages in nigrifilis are heavily pigmented.^
2. These same filaments in the new species are only one-fourth of the length of the illicium from basal bone-articulation to distal end of the bulb; in nigrifilis they are about one-half this length or relatively twice as long.
3. In alhifilosa the stalk of the illicium is heavily pigmented and this pigment runs out distally so far that it covers one-half of the terminal expansion containing the illicial bulb. In nigrifilis neither the illicial stalk nor its terminal expansion are so pigmented (Regan and Trewa- vas, 1932, Fig. 92).
4. The compressed escal appendage in alhifilosa is somewhat longer relative to the length of the illicium and it is rather more lightly pig- mented than it is in nigrifilis.
5. The illicium is only 40% rather than 50% of the standard length in the new species.
6. In the present specimen the illicium distal to the basal bone- articulation is only 2.5 times the length of the exserted part of the pterygiophore whereas in nigrifilis it is six times this length (see, however, footnote 2, p. 85).
7. The cleft of the mouth in alhifilosa does not extend posteriorly to the level of the eye whereas it extends behind the eye in Regan and Trewavas' (1932) specimen of nigrifilis.
' It is worth pointing out that the type specimen of D. alhifilosa was so little damaged when it arrived on deck that it lived for a short time in chilled sea water (see preceding section, p. 71); presumably then such an absence of pigment as mentioned above may well be considered natural rather than due to damage in capturing the specimen.
waterman: deep-sea angler-fishes 93
8. The dentition of the two species would appear to be significantly different; albifilosa has 60% more teeth in the upper jaws, and 30% more in the lower than nigrifilis. This fact is difficult to evaluate since we do not as yet have any data on the individual or ontogenetic differ- ences in the teeth of ceratioids.'
9. The only known specimen of nigrifilis is from the China Sea (19°18.5'N.,120°13'E.) whereas albifilosa is from the western Atlantic.
The above differences which distinguish the new species from its nearest relative are thus seen to be rather small, and when considered separately some of them are of questionable importance. Nevertheless, they are of the same order of magnitude as those which have been used to establish a great many of the species of this genus (compare, for example, D. ptilohis, D. viultifilis, and D. claviger Regan and Trewavas, 1932, p. 73). From the ninety-three specimens of Dolo- pichthys which have been taken forty-six species have been described ; twenty-nine (63%) of them are monotypic. The distinctions between the various members of the genus are, however, of two principal orders of magnitude so that Regan and Trewavas (1932) have distinguished between these by grouping certain of the species differing inter se by such characters as minor variations in the escal pattern into five sub- genera which are characterized by differences of a more fundamental nature.- In spite of the small amount of material available for com- parative study and in spite of the slight variations which have been used in establishing many species, from their wide experience Regan and Trewavas (1932, p. 71) feel that most of the species will prove in the future to be valid.
In view of these facts the establishment of a new species for the present specimen would seem to be quite in keeping with the precedent for the genus.
' See Fraser-Brunner, 1935, for discussion of evidence for possible ontogenetic changes in the dentition of Gigantactis.
- Parr (1934, p. 4 Iff.) in his revision of the genus Linophryne has accomplished the same sort of distinction between the relative importance of characters by the alternative method of setting up sub-species.
I
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BIBLIOGRAPHY
BOULENGER, G. A.
1904. A synopsis of the suborders and families of Teleostean Fishes. Ann. Mag. Nat. Hist., Ser. 7, Vol. 13, pp. 161-90.
Brauer, a.
1902. Diagnosen von neuen Tiefseefischen, welche von der Valdivia-Expedition gesammelt sind. Zool. Anz., Bd. 25, S. 277-98.
Brauer, a.
1906. Die Tiefseefische. I. Systematischer Teil. Wiss. Erg. Deut. Tiefsee- Exp. . . . "Valdivia," Bd. 15, Lief. 1, S. 1-420.
Fraser-Brunner, a.
1935. New or rare fishes from the Irish Atlantic slope. Proc. Roy. Irish Acad., Vol. 42B, pp. 319-26.
Gregory, W. K. and G. M. Conrad
1936. The evolution of the pediculate fishes. Amer. Nat., Vol. 70, pp. 193-208.
Parr, A. E.
1934. Report on experimental use of a triangular trawl for bathypelagic collecting. Bull. Bing. Oceanogr. Coll., Vol. 4, Art. 6, pp. 1-59.
Regan, C. T.
1925. New Ceratioid fishes from the N. Atlantic, the Caribbean Sea, and the Gulf of Panama, collected by the "Dana." Ann. Mag. Nat. Hist., Ser. 9, Vol. 15, pp. 561-7.
1926. The Pediculate Fishes of the Suborder Ceratioidea. Danish "Dana" Exp., 1920-22, Report No. 2, pp. 1-45.
Regan, C. T. and E. Trewavas
1932. Deep-sea angler-fishes (Ceratioidea). Carlsberg Found. Oceanogr. Exp., 1928-30, Report No. 2, pp. 1-113.
i
Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
I Vol. LXXXV, No. 4 \
THE LEPIDOPTERA OF BARRO COLORADO ISLAND, PANAMA
By William T. M. Forbes Cornell University, Ithaca, New York
With Eight Plates
CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
August, 1939
PUBLICATIONS OF THE
MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD COLLEGE
There have been published of the Bulletin, Vols. I to LXXXV, No. 1, 2 & 3; of the Memoirs, Vol. 1 to LIV, No. 1, 2, 3 & 4, and Vol. LV.
The Bulletin and Memoirs are devoted to the publication of original work by the Officers of the Museum, of investigations carried on by students and others in the different Laboratories of Natural History, and of work by specialists based upon the Museum Collections and Exploration.
These publications are issued in numbers at irregular intervals. Each number of the Bulletin and of the Memoirs is sold separately. A price list of the publications of the Museum will be sent on application to the Director of the Museum of Comparative Zoology, Cambridge, Massachusetts.
Bulletin of the Museum of Comparative Zoology
AT HARVARD COLLEGE
Vol. LXXXV, No. 4
THE LEPIDOPTERA OF BARRO COLORADO ISLAND, PANAMA
By William T. M. Forbes Cornell University, Ithaca, New York
With Eight Plates
CAMBRIDGE, MASS., U. S. A.
PRINTED FOR THE MUSEUM
August, 1939
/^K^^ Zoology *'/^,
AUG 31 J939 "
No. 4. — The Lepidoptera of Barro Colorado Island, Panama
By William T. M. Forbes /^seumorj^^
Cornell University, Ithaca, New Yorkf >1^g **'**»> "♦^^
TABLE OF CONTENTS ^*^1^
Page Family EUCHROMIIDAE 99
Subfamily EUCHROMIINAE 106
Pompiliodes 106
Homoeoeera 106
Autochloris 107
Sarosa 107
Gymnelia 108
Phoenicoprocta 110
Pheia Ill
Loxophlebia 112
Mesothen 112
Chrostosoma 113
Cosmosoma 114
Rhynchopyga 119
Saurita 120
Psoloptera 123
Dycladia 124
Macrocneme 124
Calonotos 134
Subfamily CTENUCHINAE 135
Dinia 135
Trichura 135
Marecidia 136
Argyroeides r 137
Hypocladia 137
Eumenogaster 138
Metastatia 138
Syntrichura 139
Chrysostola 139
Ecdemus 140
Pterygopterus 140
Episcepsis 141
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Page
Androcharta 144
Amycles 144
Leucopleura 145
Atyphopsis 145
Napata 145
Aclytia 146
'Delphyre 148
Heliura 151
Eucereon 152
Correbia 159
Correbidia 160
Hyaleucerea 161
Family NOLIDAE 162
Roeselia 165
Nola 168
Family ARCTIIDAE 168
Subfamily LITHOSIINAE 169
Hypareva 172
Agylla 173
Apistosia 175
Antona 176
Thyone 176
Odozana 177
Phaulosia 178
Prepiella 179
Callisthenia 180
mice 181
Nodozana 182
Gaudeator 182
Lycomorphodes 184
Talara 184
Dolichesia 186
^ Clemensia 187
Diarhabdosia 187
Paraprepia 188
Metallosia 188
forbes: lepidoptera of barro colorado v
Page
Subfamily ARCTIINAE 189
Robinsonia .■ 192
Idalus 193
Phaeomolis 194
Eupseudosoma 194
Premolis 195
Neritos 196
Neonerita . 197
Parevia 198
Glaucostola 198
Automolis 199
Castrica 203
Melese 204
Ammalo 205
Baritius 206
Carathis 207
Ochrodota 207
Tricypha 207
Pelochyta 208
Elysius 208
Thalesa 209
Halysidota 209
Purius 214
Calidota 214
Tanada 215
Metaxanthia 215
Agoraea 216
Bituryx 216
Belemnia 218
Diospage 218
Belemniastis 219
Virbia 220
Ecpantheria 221
Apantesis 222
Family PERICOPIDAE 223
Eucyane 224
Pericopis 224
Hyalurga 226
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Page
Family AGARISTIDAE 230
Darceta 230
Family LYMANTRIIDAE 231
Caviria 232
Thagona 233
Sarsina 234
Sta?therinia 234
Desmoloma 235
Family NOTODOXTIDAE 235
Subfamily MELALOPHINAE 246
Rosema 246
Subfamily NOTODONTINAE 247
Calledema 247
Elymiotis 249
Pentobesa 250
Nystalea 251
Phedosia 254
Proelymiotis 255
Bardaxima 256
Lepasta 256
Strophocerus 258
Tachuda , 259
Antiopha 262
Marthula 263
Crinodes 263
Dasylophia 265
Betola 268
Didugua 268
Lusura 268
Lirimiris 269
Canodia 270
Phyllopalpia 270
Peroara 271
Apela 271
Dicentria 281
Astapa 282
FORBES: LEPIDOPTERA OF BARRO COLORADO VII
PAGE
Naprepa 282
Kalkoma 283
Malocainpa 284
Farigia 286
Disphragis 286
Talmeca 293
Rifargia 294
Meragisa 297
Magava 297
Phastia 298
Drugera 298
Rincodes 299
Blera • 299
Rhuda 300
Anita 301
Colax 301
Hemiceras 302
Hapigia 314
Rhapigia 316
Chliara 317
Antsea 317
Subfamily CERURINAE 318
Cerura 318
Family DIOPTIDAE 318
Phseochlfena 319
Momonipta 319
Scotura 320
Actea 320
Josia 320
Getta 322
I
A. Euchromiidoe, Nolidoe, Arctiidoe, Pericopidoe
The material of this report should give a fair sample of the fauna of Barro Colorado Island, and is presented to put the new data on record, but also to serve as a basis for other studies, biological and the like, which may be made there. It is based on collections made all through the year except for a short gap in the latter part of August and September, but more systematically in November to February, 1934-35, when Dr. Marston Bates was collecting continuously and in- tensively, and in February and March of 1936, when a group of the staff of the American Museum of Natural History worked there. While many more species will certainly be taken, especially those whose main flight periods come in the summer, it is probable that all the really dominant species are included, and that except for mid- summer, few of the more conspicuous forms are omitted.
The great mass of the material is in the Museum of Comparative Zoology. The American Museum has the lot collected in Feb. -Mar., 1936, by F. E. Lutz, W. J. Gertsch and Wm. C. Wood, and a few col- lected by C. H. Curran in 1935, and Cornell a few collected by J. C. Bradley in Mar.-iVpr. 1924. The principal collectors for the Museum of Comparative Zoology were Dr. Bates, Mr. A. Friedman in the early summer of 1935, Graham Fairchild in the summer of 1923, and Nathan Banks in the summer of 1924; the last two contributing relatively few specimens but giving us our only knowledge of the summer fauna.
Another lot that has been mentioned and included in the keys is that taken by Mr. August Busck in the part of the Canal Zone which has since been flooded, in 1911-12. So far as -this fauna survives it must have taken refuge on Barro Colorado Island, and a few more limited areas. The records were published by Dyar\ and a few more have been added which were identified or described more recently from the same lot, in most cases by Dr. W'illiam Schaus.
For the sake of compactness, collectors' names will be cited specifi- cally only in records of unusual character, and all records not other- wise credited are to be assumed as referring to Barro Colorado material, now in the Museum of Comparative Zoology. Duplicates
iProc. U. S. Nat. Mus. xlvii, 1-67 (Busck), 139-350 (Dyar) ; 1914.
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of most of the species will be deposited in the Cornell University Collection.
The four families discussed here form a homogeneous group, closely related to the Noctuidse, with which they share the "quadrifid" vena- tion and the thoracic tympanum. Larval characters would indicate that the Nolidse are directly derived from the Noctuid?e (Sarrothrip- inse), but that the others are of a parallel stock arising from the Hypsidse.
The get-up of the paper is designed to be convenient for those in- tending to identify material as a basis for biological studies; so brief keys are included of the species actually taken on the Island, and those included in the National Museum survey; and the bibliographic citations emphasize the published figures.
The following are the principal works on the area :
Dbtjce, H.: Biologia Centfali-Americana, Lepidoptera-Heterocera. 2 vols, text and one of plates (abbreviated in the citations of figures as "Biol." with plate and figure numbers).
Hampson, G. F. : Catalogue of the Lepidoptera PhalsentB, i and Supplement i (Euchromiidse as Syntomida? or Amatida;), ii and suppl. i (Nolidse; Litho- siinse), iii and suppl. ii (Arctiinaj); (abbreviated "Hamps." and "Hamps. Suppl." with plate and figure).
Seitz, a.: Macrolepidoptera of the World or Grosschmetterlinge der Erde; vi, Euchromiida?, NoHdae and Lithosiinte by M. Draudt, Arctiinaj by Seitz, Pericopida; by M. Hering; (abbreviated "Seitz" with plate, line, and figure counting from the left).
RoTH.scHiLD, Walter: Novitates Zoological xvii, 1-85, 113-171, 1910 (giving fuller distributions of many species).
Dyar, H. G.: Proc. U. S. National Museum xlvii, 139, 1914, (giving further Canal Zone data for many species). ^
In the case of the first three works the numerous figures are cited very briefly; — other figures referred to are cited sufficiently fully to lead directly to their location. In general the figures in Seitz are copies of the others, and frequently have lost something in copying. It also figures a few species described since Hampson's supplements.
Where descriptions of early stages have been noted they are cited in the bibliographies, and summarized in the text, but otherwise the
• Rothschild has also published a series of papers on these families supplementary to Hamp- son's Catalogues. The principal ones are Nov. Zool. xvi, 21-55, 268-299, pis. 4-7; xvii, 172-188, 429-445, 504-506, pis. 11-14; xviii, 24-45, 154-158, pis. 3-6; xix, 151-186, 212-246, pis. 3-5; XX, 192-226, pis. 13, 14.
FORBES: LEPIDOPTERA OF BARRO COLORADO 99
bibliography is cut to the original description, reference to important or convenient figures, and new or obscure synonymy. A somewhat more complete bibliography will be found in the Lepidopterorum Catalogus, fascicles vii (Syntomidse, i.e. Euchromiidte, by Zerny, 1912), xxii (Arctiinse by Strand, 1919), xxiv (Nolidie by Strand, 1920), xxvi (Lithosiinae by Strand, 1922) and xlv (Pericopidse by Bryk, 1931).
Dates for the commoner species are presented in tabular form at the heads of the families Euchromiidse and Arctiidae, — those of rarer forms are included in the text.
EUCHROMIID.E
(Syntomidse; Amatidse; Zygpenidse of earlier authors)
Head with strong palpi and tongue and conspicuous ocelli; thorax with tympanum below base of hind wing (vestigial in the Old World types), protected by a hood developed from the first segment of the abdomen above the spiracle; fore wing with apparently 4-branched Cu (except a few species which have lost M2); hind wing relatively small, Sc normally absent, in some primitive species visible as a short spur from upper side of cell beyond middle. Larva with a single large subdorsal wart each on meso- and metathorax; with feathery tufted hair and usually pencils, which are most often lateral on first and 8th segments of abdomen. Pupa fusiform, without cremaster or flange- plates on sides of abdomen, with antennae and tongue extending to tips of wings; glossy.
The absence of Sc will separate this family as a rule from the Arctii- dse, but in Eucereon, Episcepsis, Aclytia and Hyaleucerea there are species with Sc preserved, while many Phegopterine Arctiidae have Sc more or less aborted, and a few have lost it entirely. The Euchromii- dae, unlike the Arctiidae, are almost always heavily pigmented forms with brown or black veins, while most of the Arctiidae are lightly pig- mented night-flyers. The Belemnia group would be an exception but should probably be transferred to the Euchromiidae.
I should divide the family into three subfamilies. The Amatinoe are limited to the Old World, and all save a few primitive African types (Pseudapiconoma, Melisa and Phaeosphecia) have fused R and Mi, so that the hind wing has only a single vein from the upper angle of the cell. In the Euchromiinas M2 of the hind wing is rudimentary or absent, being represented by a faint vein or line of scales from the angle of the discocellular, and Cui and Cu2 are stalked or united; while in the Ctenu-
100 bulletin: museum of comparative zoology
chinae M2 is fully developed and curved, arising from below the angle of the discocellulars, and Cui and Cu2 are widely separated except in the Horama group. A few aberrant genera also have M2 obsolescent from the angle but Cu2 well separated. I believe these are aberrant Ctenu- chinte, but none have yet been taken on Barro Colorado Id. The two latter subfamilies are New-world, except the single genus Euchromia; and the Nearctic species are all Ctenuchinse, one or two Euchromiinae reaching north to Georgia.
The diagram opposite records the dates of capture of the commoner species, as represented in the collections of the Museum of Compara- tive Zoology. Each column represent a quarter of a month. Where a single specimen was taken it is indicated by an open circle (o), if more than one by a solid spot (•). In the case of Fairchild's specimens, which were not individually dated, records are indicated by a line covering the proper period, and the number of specimens. Except for such of these as may have been taken in early August, August and September are vacant. Dyar (I.e.) gives further regional dates.
Key to Genera of Euchromiidae
1. Hind wing with Cui and Cu2 stalked or united, in the latter case the
hind wing with only three dorsal veins, and the two lower (which are M3 and Cui+2) arising closer together than to M2, or with only two dorsal veins, M2 being a rudimentary fold or line of scales extending from the angle of the discocellular {Euchro-
miincB and Horama group)^ 2
Hind wing with Cu2 widely separated; M2 and M3 sometimes united, in which case the two most dorsal veins (Cui and CU2) are well separated and the upper (M2+3 and Cui) are close together or stalked, never with only two dorsal veins {Ctenuchince) 22
2. Upper of the three veins (M2) separated by a distinct space from
the stem of the other two at origin 3
Upper of the three veins (normally M3) connate or short-stalked with the stem of the other two at origin 4
3. Head and prothorax densely hairy Amycles
Head and prothorax smoothly scaled Ceramidia
• In the Euchromiinae there are only 3 veins by the obsolescence of M2; in the members of the Horama group which have been reported from the Zone there are only 3 veins by the complete union of Cui and Cuj, in Horama itself, which is regional, Cui and Cuj are forked toward the margin.
FORBES: LEPIDOPTERA OF BARRO COLORADO
101
|
n o |
O N. CO uo lO CO TfCCcC |
CO O CO CO C^l C^l C^ O — Ol |
1^ O CO lO •-< O f 01 Tf o |
Ol 1^ lO O 05 Ol — -J5 — |
CO O C2 o o |
CO coo COIN ^ |
OS ceo-* ■*(N.-i |
|
Dec. 12 3 4 |
•om ••• ©•••o |
©••o •• • ••• |
©• • • • ^00 |
• o o o • • |
•©•• • o • |
• • • ©• o ••o |
• • • • •ooo •©•o |
|
Nov. 12 3 4 |
•oo^ • • • |
©•• o • •o o |
•©• • o •o o |
•o o ©• |
oo^ oo o o |
•oo oo • oo |
••© •© ©•© •©© |
|
Oct. 12 3 4 |
om o •o o |
•o o |
o • • • |
• |
• o o |
••• o ©© © |
|
|
Aug. 12 3 4 |
. . |
. . |
|||||
|
July 12 3 4 |
-r- S |
* |
^'p< —' |
IM rt rt |
•• |
M |
•—1 1—1 |
|
June 12 4 4 |
O • o o |
• |
o o o |
o o o |
O oo |
o |
|
|
May 12 3 4 |
oo o o o |
o |
o o o |
O OOO |
o oo |
||
|
Apr. 12 3 4 |
o o om m •• |
o •O O |
o o oo •o o |
• o • o •o |
o o •o |
o oo |
o o |
|
Mar. 12 3 4 |
• ©••• ©©••^ ••• |
OO^ OO • |
o • •©• • o |
o o • •o • |
©• oo^ ©• o |
o o • o o |
|
|
Feb. 12 3 4 |
••••o •©•• ••••o ••••o |
O • •• o ••• • oo^ o |
• •oo • ••©©• ••o • |
o • • • o •••oo ©••o^ |
• o o ©• ©• • |
o • •o ©•o |
m ©•oo ••• |
|
Jan. 12 3 4 |
••••o oo • o oo^ |
•©•o^ • o ©•• • |
•• o^ • • ©• oo •• |
• oo •o o oo o o |
©• •• • o o o oo |
©•• o •o |
•o© •© © |
|
Month Week |
p. aliena II. stictosoma G. salvini G. colona P. paucipuncta |
M. pyrrha M. ethela C. echenius C. tabascensis C. saron |
C. metallescena C. hercynacula S. clusia S. nox S. phoenicosticta |
S. anselma S. afflicta P. thoracica D. correbioides M. auripes |
M. cyllarus C. tiburtus M. achrysa H. re.stricta Ec. obscurata |
D. rubricincta Eu. obscurum C. affinis |
C. e. germana C. e. costinotata C. e. elegans C. e. striata |
|
< 2; |
102 bulletin: museum of comparative zoology
4. Fore wing with Ri and 2 stalked (normally arising out of the base
of R3-5) 5
Fore'wing with R2 stalked with R 3-5, Ri separate 6
R2 arising separately from cell 21
5. Wings with transparent spots or areas Leucopleura
Wings opaque and evenly scaled Psoloptera
6. Abdomen with small divergent sublateral tufts on last segment;
the Barro Colorado species markedly constricted at third seg- ment Phwnicoprocta
Abdomen without lateral tufts on last segment, the abdomen con- stricted at second segment if at all 7
7. Second segment of abdomen narrowed by a slight notch in front,
abruptly widened behind, wasp-like Pompiliodes
Second segment of abdomen not constricted 8
8. Antenna with middle part of shaft considerably widened as com-
pared with its base, especially in female 9
Antenna with shaft not widened, or in female with middle of shaft a little wider than extreme base 13
9. Disc of thorax with considerable long hair Homoeocera
Thorax with long hair only on edges of tegulse etc., mostly close- scaled 10
10. Hind tibiiie toward end and tarsi fringed with long scales, at least
above; blade-like Macrocneme
Hind tibiae with the usual close scaling 11
11. Hind wing with lower part of cell very short, reaching only about
1/3 way to apex of wing; wings transparent Isanthreme
Hind wing with lower side of cell longer, reaching almost half length of wing 12
12. Cui of fore wing arising almost half way between M3 and Cu2
Dycladia Cui of fore wing arising about 3^ as far from M3 as from Cu2
Autochloris (Bombyliodes)
13. Fore wing with M2 leaving cell well above origin of M3 14
Mo and 3 closely approximate or connate at origin 15
14. Hind wing with lower discocellular very short, obsolescent
LoxophJcbia Hind wing with lower discocellular a fifth as long as middle one
Mesothen
15. Hind wing with lower discocellular obsolescent, about l/lO as long
as middle; lower side of cell less than 3^ length of wing. . . Pheia
FORBES: LEPIDOPTERA OF BARRO COLORADO 103
Hind wing with lower discocellular about 1/6 as long as middle or longer; cell more than J^ length of wing 16
16. Fore wing with Ri stalked on R2-5; cell of hind wing moderate with
lower discocellular moderately long, and M3 and Cui moder- ately stalked Chrostosoma
Fore wing with Ri arising separately from cell 17
17. Hind wing with M.3 and Cui strongly stalked, l/3 or more of the
distance from end of cell to margin 18
Hind wing with M3 and Cui connate or barely stalked . . . Saurita
18. Thorax densely hairy, except patches on the collar and sometimes
shoulders, tv^hich are smoothly scaled Sarosa
Thorax with at least large areas on tegulpe and disc closely scaled as well as collar, frequently with only fringes of long hair on tegulfe etc 19
19. Wings solidly scaled, at most with small transparent streaks on
basal part of fore wing and base and disc of hind wing. Rhynchopyga Wings largely transparent, usually less elongate 20
20. Outer margin of hind wing evenly rounded or nearly so (sometimes
somewhat lobed in small species with dominantly yellow bodies).
Cosmosoma Outer margin of hind wing with anal lobe separated by a distinct notch; abdomen dominantly dark with blue spots; thorax nor- mally more hairy Gymnclia
21. Fore wing with M2and M3 closely parallel about J^way to margin,
then divergent; hind tibia but slightly flattened and not fringed
Calonotos
Fore wing with M2 and M3 immediately divergent; hind tibije and
tarsi with strong fringes of hair-scales above . . Macrocneme (part)
22. Both wings with a dorsal vein absent (M2 and M3 united) ; small
wasp-like species with transparent wings Syntrichura
Fore wings with all dorsal veins present 23
23. Hind wing with a dorsal veing absent 24
Both wings with all veins present 27
24. Wings opaque or with small translucent spots; abdomen simple;
hind wing with cell normal and the interspace between M2 + 3
and Cui twice as long as wide Delphyre
Wings broadly translucent or transparent; abdomen with lateral tufts on terminal segment 25
25. Wings perfectly hyaline (with black markings) ; cell of hind wing
normal, abdomen with a pair of lateral tufts on last segment
Metastatia
104 bulletin: museum of comparative zoology
Cell of hind wing very long, the interspace between veins M2 + 3 and Cui about as wide as long 26
26. Wings hyaline; abdomen constricted on third segment but con-
tinuing narrow, and not specially marked there; abdomen with
only anal tufts Chrysostola
Wings translucent; abdomen strongly constricted at third segment and with white spots, making it seem even more slender; with lateral as well as anal tufts Eumenogaster
27. Abdomen more or less constricted at second segment 28
Abdomen not constricted, or slightly constricted at third segment {Marccidia) 30
28. Abdomen strongly constricted, wasp-like; the anal area of hind
wing reduced to a hairy lobe 29
Abdomen moderately constricted ; anal lobe normal i^thria
29. Shaft of antenna thickened and roughly scaled above near middle;
male abdomen frequently with a long terminal filament
Trichura Shaft of antenna smooth and slender Argyroeides
30. Antenna with middle part of shaft fringed with long hair; wings
hyaline Dinia
Antenna with shaft smoothly scaled; wings normally fully scaled 31
31. Palpi upturned, with third joint also erect, continuing the general
line of the palpus 32
Palpi upturned or oblique, but with the third joint porrect, fre- quently long 43
32. Hind wing with M2 arising from close to angle of cell or stalked
withMa 33
Hind wing with M2 well separated from M3 38
33. Fore wing with Cui concave above, separate from M3 at origin and
strongly convergent to Cu2 at margin ; hind wing with M3 and
Cui stalked Mydromera
Fore wing with Cui and Cu2 normally divergent to margin ... 34
34. Both wings with M2 and M3 distinctly stalked; wings translucent
AtyijhoiJsis
Fore wing with Mo and M3 separate, hind wing with them separate
or shortly stalked ; wings normally opaque 35
35. Fore wing with R2 stalked on R3-5 36
Fore wing with R2 free Hyaleucerea
36. Basal segments of abdomen dorsally or subdorsally clothed with
rough hair Eucereon
Abdomen smoothly scaled to base 37
FORBES: LEPIDOPTERA OF BARRO COLORADO 105
37. Hind wing of normal size; tympanic hoods conspicuous above
Napata
Hind wing relatively small, more or less aborted in male; tympanic
hoods inconspicuous Androcharta
38. Hind wing with M3 and Cui stalked 39
Hind wing with M3 and Cui arising separately from cell 40
39. Fore wing with Ro separate; Mi connate with R3-5 Aclytia
Fore wing with R2 stalked on R3-5; Mi arising distinctly separated
Heliura
40. Abdomen dorsally or subdorsally clothed with rough hair toward
base 41
Abdomen smoothly scaled to base Hypocladia
41. Fore wing with M2 and M3 approximated for some distance from
origin Ptergopterus
Fore wing with Mo and M3 divergent from origin 42
42. Hind wing with M3 and Cui widely separated, with the trans-
parent area sharply bounded ; typically with R and Mi stalked, and with R2 free in fore wing, but the present species with R
and Ml of hind wing and R2 of fore wing stalked Ecdemus
Hind wing with M3 and Cui connate, the transparent area vague and partly scaled over; R2 always stalked; R and Mi of hind wing connate or barely stalked Episcepsis
43. Shaft of antenna widened beyond middle; hind wing small, less
than half as long as fore wing, the anal area reduced with a single anal vein; hind tibia of male much enlarged, with sex-scaling
Marccidia
Antenna shaft normal; hind wing relatively larger, frequently large with ample anal area ; hind tibia normal 44
44. Middle and hind tibise smooth Ctenucha
Middle and hind tibia tufted at the spurs 45
45. Palpus with third segment set at an angle to second, which is
oblique, and half as long; tip of second segment tufted; fore wing with M2 and M3 long-stalked, the wing very narrow
Correbia
Palpus with third segment continuing the line of the second or with a gentle down-curve; wings moderately broad, M2 and M3 separate or only short-stalked, M3 and Cui often stalked
Correbidia
lOG bulletin: museum of comparative zoology
Subfamily EUCHROMIINAE
PoMPiLiODES Hampson
A curious feature of this family is the close parallelism of pattern in members of the two subfamilies. Thus the perfect type of wasp- mimicry is duplicated in Pseudosphex and Argyroeides, though the narrowing at the "waist" takes place in a different way; the present genus (which is a mimic of Parachartergus) is duplicated by Amycles; Macrocneme by Horama; the aberrant Chrostosomas by Metastatia and Cacostatia, Dycladia by Correbia and Correbidia and so on. In this genus the wasp-waist is evanescent, but the general appearance is perfectly wasp-like.
322^ PoMPiLiODES ALIENA Walker
Euchromia (Pampa) aliena Wlk. List Lep. Ins. B.M. i, 241, 1854. Figured: Hamps. fig. 76^; Herrich-Schseffer, Aussereur. Schm. 73 (as Glaucopis flavofascia) 80 (as Amycles (f) /.), fig. 231; Seitz 10: k3.
Blackish, with a pale patch near apex of fore wing, large in male, more or less obsolete in female. P. posfica Wlk. is merely the more southern race, with normally smaller pale patch. P. tenebrosa Wlk. differs in the mainly transparent hind wing.
Chief flight at end of Jan. (see diagram). Guatemala to Amazons.
HoMCEOCERA Felder
This genus begins the Cosmosoma complex, which extends to Saurita, probably the ancestor of the group. The genera are close and partly artificial, and the hairy vestiture is probably secondary, Sarosa, e.g. having two groups, one derived from Cosmosoma, the other from near Gymnelia. Homoeocera has a lobed anal angle like Gymnelia, from which it also derives. The venational characters vary some and the key may not allow enough for individual variation, though it has been checked.
333 HoMCEOCERA STICTOSOMA Druce
H. stictosoma Dr. Ann. Mag. Nat. Hist. (7) i, 402, 1898. Figured: Hamps, 6: 17; Seitz 11: 4.
Easily recognized by the abdominal spots; subdorsal on 1, lateral on 2, both dorsal and lateral on 3.
' The numbers attached to the species refer to the appropriate volumes of Hampson's Cata- logue of the Lepidoptera Phalsense; the simple numbers are those cited as species in vols, i-iii, those with a decimal to forms cited there as subspecies, and those with a letter to the corresponding volumes of the Supplement (i, ii).
^Figures so cited are in black and white, showing pattern and venation.
FORBES: LEPIDOPTERA OF BARRO COLORADO 107
Dominant at end of Jan. (see diagram, p. 101). Described from "Colombia" (at a date when Panama was part of Colombia).
Dyar reports Isanthrene crabroniformis Stand. (Biol. 6: 23; Seitz 9. 14), from the Canal Zone. It is yellow-hyaline, with black, yellow and blue abdomen.
AuTOCHLORis Hiibner (Bombyliodes Hampson)
Differs from Gymnelia only in the more broadened antennse. I follow Hampson in changing this name (Suppl. p. 102). It is a question of the identity of Autochloris almon (Cr.). The larva of A. almon as figured by StoU (11: 1) is mouse gray, with 4 long anterior and two posterior black pencils, tipped with white.
363 Autochloris jansonis Butler
Gymnelia jansonis Btl. Cist. Ent. i, 116, 1872. Figured: Butler, Lep. Exot. 61: 17; Seitz 11: b2.
Abdomen with a cream white bar across first segment, and orange streaks on middle segments. Three females.
May 25 (Friedman), July 12, 1924 (Banks); Nov. 9-10, (Am. Mus. Nat. Hist.). Costa Rica and Panama.
Sarosa Walker
This is a heterogeneous genus, S. innotata being a hairy variant of Cosmosoma (with the typical forms even more hairy) while lutihasis is merely a somewhat more hairy Autochloris. The female is less hairy than the male, female lutihasis being only about as hairy as males of Autochloris.
1. Disc of thorax and most of body yellow innotata
Body black with a double yellow patch at junction of thorax and abdomen lutihasis
(380.1) Sarosa innotata Draudt
Sarosa notata, ab. 1 Hamps. Cat. Lep. Phal. i, 187, 1898.
Sarosa notata, form innotata Ddt. in Seitz Macrolep. World vi, 53, 11: f5, 1915. ? Paecilosoma semirubra Ddt. in