TAXONOMY AND BIOLOGY OF THE GENUS LEBETUS (TELEOSTEI-GOBIOIDEA) BY P. J. MILLER (Department of Zoology, The University, Glasgow, W.2) Pp. 205-256 ; Plate ; 21 Text-figures BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY) ZOOLOGY Vol. 10 No. 3 LONDON : 1963 THE BULLETIN OF THE BRITISH MUSEUM (NATURAL HISTORY), instituted in 1949, is issued in five series corresponding to the Departments of the Museum, and an Historical series. Parts will appear at irregular intervals as they become ready. Volumes will contain about three or four hundred pages, and will not necessarily be completed within one calendar year. This paper is Vol. 10, No. 3, of the Zoological series. Trustees of the British Museum 1963 PRINTED BY ORDER OF THE TRUSTEES OF THE BRITISH MUSEUM Issued March 1963 Price Twenty One Shillings TAXONOMY AND BIOLOGY OF THE GENUS LEBETUS (TELEOSTEI-GOBIOIDEA) By P. J. MILLER ABSTRACT The marine gobioid genus Lebetus Winther 1877 is redefined and previous division into two species orca Collett 1874 and scorpioides Collett 1874 shown to be based on sexual dimorphism in one species, by page priority L. orca. A redescription of the species is provided including details of sensory papillae and osteology. L. orca is eastern Atlantic boreal in distribution, occurring as a small predator on coarser grounds down to 375 m. Specialization of the male genitalia and sexual dimorphism are described and the breeding season provisionally regarded as from March to August. The skeleton of Lebetus is typically gobiid, and evidence is presented for a closer relationship to the genus Buenia Iljin as represented by B. Jeffrey sii (Giinther) than to the Mediterranean Odondebuenia De Buen and Cabotichthys Whitley with which Lebetus had been previously grouped. INTRODUCTION THE genus Lebetus was founded by Winther (1877) to contain two species of gobies dredged by G. O. Sars on the coast of Norway and described by Collett (1874) as Gobius orca and G. scorpioides. These species have subsequently been regarded as among the rarest of European Atlantic fishes and, until recent years, the numbers of demersal individuals known to science have been relatively few. In the last fifteen years, protracted dredging surveys off the Isle of Man (Jones, 1951 ; Hartnoll, 1961) have yielded many examples. Their small size and occurrence on rough grounds makes collecting difficult ; in Manx waters, most specimens have been obtained by means of a scallop dredge lined with shrimp netting. Relative infrequency of capture is probably due to inadequacies of the fishing gear rather than to actual rarity, and occasional hauls providing up to five individuals suggest the presence of appreciable numbers in the area. Other long term faunistic investigations within the range of the genus have produced more or less sizeable collections of both demersal (Grieg, 1913 ; Le Danois, 1913) and planktonic stages (Petersen, 1919 ; Johansen, 1925 ; Russell, 1926-1940). My interest in these fishes was aroused firstly by the difficulty experienced in separating the two species. After endeavouring to use the diagnoses of several authors, it was found possible to achieve this only on the basis of coloration. Then, while examining the testes of various gobies, it was noticed that no female L. orca were present in a collection of twenty two dissected individuals, and only immature males among twenty eight L. scorpioides. The possibility that the two so-called species were in fact based on sexual dimorphism in a single species indicated the need for taxonomic revision. Although such a view was put forward by Smitt (1900), this was not accompanied by any evidence and appears to have been overlooked by later authors. Dissection of the extensive material from the Irish Sea presented oppor- tunities for settling the position of Lebetus among the gobioid families by osteological ZOOL. 10, 3 i4 208 P. J. MILLER study, and also for investigating various aspects of diet and reproduction. A pre- liminary account of some of this work has already been published (Miller, 19616). MATERIAL AND METHODS The following specimens have been available for examination : Isle of Man : 33 $<$, standard lengths 15 -75-30-0 mm., including BMNH 1961 . 10 . 17 . i. 3 5 an( i one m PEM, and 27 ?, 14-5-29-0 mm., inc. BMNH 1961 . 10 . 17 . 2, 4, and one in PEM. Other British localities : 3 $, 12-0-30-0 mm., BMNH 88 . 3 . 22 . 9-10 part and BMNH 1903.4.14.4-7 part, and 2 $$, 10-5 and 17-0 mm., BMNH 1903.4.14.4-7 part. Scandinavia: 13 <$<$, 13-5-26-0 mm.,ZMO J3999 (type of Gobius orca), J40OO, J4OOI, J4023 ; ZMB 771, 1887, 1893, 1966, 4174, 4175, 5294 ; ZMC 89, 91 ; and 4 ??, 21-75-29-0 mm., ZMO J402O, J4O2I (types of G. scorpioides) ; ZMB 536, 2009. Abbreviations for museums are as follows : BMNH British Museum (Natural History). PEM Port Erin Marine Biological Station Museum. ZMB Zoologisk Museum, Universitetet i Bergen. ZMC Universitetets Zoologisk Museum, Copenhagen. ZMO Zoologisk Museum, Universitetet i Oslo. The specimens were viewed by means of low power binocular and dissecting micro- scopes. With suitable illumination the sensory papillae were readily seen without staining after careful removal of mucus from the head and body surfaces. The skeleton was investigated by dissection of alizarin stained whole-mount preparations. THE GENUS LEBETUS WINTHER 1877 Lebetus Winther, 1877: 49 (orthotype Gobius scorpioides Collett 1874); Smitt, 1900: 554; Jordan, 1919 : 392 ; 1923 : 225 ; Duncker, 1928 : 140 ; Iljin, 1930 : 55 ; De Buen, I93oa : 123 ; 19306 : 5, 21 ; Koumans, 1931 : 43, 46 ; Whitley, 1931 : 155 ; De Buen, 1931 : 50, 54, 61 ; 1940 : 3 ; 1951 : 56, 57 ; Norman, MS : 413. Lebistes Jordan, 1920 : 487 ; 1923 : 226 (non Filippi, 1862 : 69). Butigobius Whitley, 1930 : 123. Body moderately elongate, subcylindrical, with 25-29 ctenoid scales in lateral series. Head, nape, and back naked to origin of second dorsal fin, lacking skin folds or barbels. Postorbital length not more than half length of head. Anterior nostril a short tube. Opercle without scales. Preopercle and lower jaw unarmed. Branchi- ostegous membranes attached to sides of isthmus, but not fused across isthmus. Jaws oblique, subequal, maxillae ensheathed and not elongate. Teeth of both jaws in several rows, simple, erect. Tongue truncate to very weakly emarginate, free from floor of mouth anteriorly. Pelvic fins united along entire length, without anterior membrane (frenum) and with fourth branched ray somewhat produced. Pectoral girdle lacking flaps on anterior edge ; uppermost rays of pectoral fin not free from membrane. Dorsal fins separate, first dorsal with 6 rays, not pungent. Caudal fin rounded, not much longer than head. Cephalic canals absent. Sensory papillae prominent but reduced in number. Two pairs of interorbital papillae (/>). Cheek with only suborbital row a, a short longi- TAXONOMY AND BIOLOGY OF THE GENUS LEBETUS 209 tudinal row c, and an intermediate papilla ; row d without posterior longitudinal section. Anterior dorsal (occipital) series (excluding h) in two groups of a few papillae. Opercular transverse row ot in two sections. Preoperculo-mandibular series in several parts. In compiling his work on the genera of fishes, Jordan (1919, 1920, 1923) includes, together with Lebetus Winther 1877, the genus Lebistes Smitt 1899. Since Jordan gives " L. scorpioides Smitt " as the orthotype of this genus, the latter, as noted by Koumans (1931), is clearly an erroneous reference to Lebetus for which Smitt (actually 1900, not 1899) indicated Gobius scorpioides Collett as the type. The generic name Lebistes is preoccupied by Lebistes Filippi 1862 among the cyprinodont teleosts and, to replace this name within Jordan's list of gobiid genera, Whitley (1930) proposed Butigobius without consulting Smitt 's paper. In a later account, Whitley (1931) unravels this confused story. THE SPECIES OF THE GENUS The present section embodies a review of the various characters proposed by earlier authors as being of value in the diagnosis of L. orca and L. scorpioides. Coloration. The two species are reputed to differ in both body and fin coloration. The body of Gobius orca was described by Collett (1874, 18750) as uniformly greyish except for darker areas between the second dorsal and anal fins, and at the root of the caudal fin, while that of G. scorpioides was said to possess four or five dark brown vertical bands across the sides. After examination of a living scorpioides, Winther (1877) remarked on the colourless caudal peduncle, and both Holt & Byrne (1903) and Le Danois (1913) emphasized the importance of this pale band, sharply demar- cated anteriorly, as a means for the identification of L. scorpioides. In the original description, Collett (1874) mentioned that the second dorsal fin of G. orca had an indistinct white band in its outer half whereas the same fin in G. scorpioides showed only ill-defined banding. Alternate dark brown and snow white bands were found across the second dorsal fin of a later specimen of G. orca (Collett, 1885). Holt & Byrne (1903) regarded a superior black band to this fin as characteristic of G. orca and in their text referred to a row of white spots immediately below the dark peri- phery. These authors described and illustrated the second dorsal fin in G. scorpioides as banded with red, the markings in some cases being flanked by dark edging. With the collection of Lebetus amassed for this work, it has been possible to segre- gate the individuals into two groups on the basis of differences in the coloration of the second dorsal fin. In the first of these, termed the orca group, the second dorsal fin has a thick black superior edge, below which occur white areas and wide oblique bands, yellow or ochre in life but grey in preserved material. Within the scorpioides group, the black edge to the second dorsal fin is present but much thinner and less intense than in the orca examples. The rest of the fin bears merely four narrow double lines of melanophores, enclosing in the living fish thin red or ferruginous striae, and separated by hyaline fin membrane. Among living scorpioides, madder brown lateral marks are pronounced on the body which, in orca fishes, is more uniform grey or. yellowish grey although comparable markings may be quite prominent. A pale band on the caudal peduncle is more clearly defined in the scorpioides group, especially 2io P. J. MILLER when preserved, but living orca show this feature to a noticeable degree (cf. PI. I and Text-fig. 3) and it is shown in Collett's illustrations of Gobius orca (i875a, 1885). TABLE I. Relation of Coloration Groups to Sex and Maturity Stage No. of males Mature and No. of females Group developing Immature all stages (a) Dissected orca 21 i scorpioides . 6 22 (b) Not dissected orca . . 16 scorpioides . 5 n The relation of these two colour patterns to sex and to gonad maturation is indicated in Table la, based on fifty Manx specimens whose sex and sexual development were confirmed by dissection. The sex and approximate maturity stage in a further thirty two British and Scandinavian examples have been determined by inspection of the genital papilla (Text-fig. 17), and these results are given in Table Ib. It is apparent that mature or developing male scorpioides do not occur, and that female orca are unknown at any stage of maturation. This distribution strongly implies that the orca coloration is the male livery of a species whose females and immature males bear the markings of scorpioides. Incipient development of the orca pattern in the second dorsal fin can, in fact, be noted among some of the immature males in the scorpioides group, which usually exhibit a pronounced black spot on the distal part of the first two interradial membranes of the second dorsal fin (see Le Danois, 1910, fig. 4). The above hypothesis is further supported by the range in standard length found in the three categories shown in Table II. The examples of orca are from 18-0-30-0 mm. TABLE II. Standard Length Frequency in Coloration Groups, Expressed in 2 mm. Length Groups Standard length ( Coloration group orca . cJ scorpioides 9 scorpioides in length while male scorpioides do not exceed 23-0 mm. Taning (1940) has reported juvenile Lebetus orca of 5-15 mm., but relied for the identification on meristic characters whose validity is doubtful. Female scorpioides attain about the same maximum length as that for orca. The first dorsal fin in the orca individuals is a uniform yellowish grey edged with white, but in scorpioides is usually banded with a dark spot on the interradial mem- brane between the fifth and sixth rays. Confluence of these bands frequently occurs, and the spot may not be very distinct. This scorpioides group pattern is stated by TAXONOMY AND BIOLOGY OF THE GENUS LEBETUS an Holt & Byrne (1903) to be found only in the breeding male of " Gobius scorpioides ". Fage (1918) regarded the presence of a black spot on the first dorsal fin of a 12-5 mm. example from the Dana collections as a sign of precocious development of supposed male coloration. However, in the material under consideration, the spot is well represented in adult females and among male Lebetus occurs only in the immature where it is not uncommonly absent. The single female G. orca mentioned in the literature was taken by Patience (1906) in the Firth of Clyde. This specimen has not been traced. The diagnosis of G. orca given by Patience was not derived from his own examples but seems to be a translation from Collett (1896) and could apply to individuals of either the orca or scorpioides group. TABLE III. Meristic Characters Cited for Lebetus orca and L. scorpioides No. of fin rays VI VI VI VI VI V-VI VI 9 9 1/9 1/8-9 8-9 1/7-8 1/8-9 8 7-8 1/6 1/6-9 6-8 1/6-7 1/6-7 18-20 18-20 18-20 X/I2/X 12 X/I2/X No. of Scales in 1.1. 28 28 24 25 c. 24 25-28 25-26 24-28 Authority Collett, 1874 Collett, 18750 Collett, 1885 Lilljeborg, 1884 Smitt, 1892 Collett, 1896 Holt & Byrne, 1903 Duncker, 1928 De Buen, 19300, b Taning, 1940 Collett, 1874 Collett, 18750 Winther, 1877 Lilljeborg, 1884 Smitt, 1892 Holt & Byrne, 1903 Le Danois, 1910 Le Danois, 1913 Fage, 1918 Petersen, 1919 Duncker, 1928 De Buen, I93oa, b, 1932 1 Scales in 1.1. c. 26-c. 28 (Grieg, 1913). z Scales in 1.1. 26, T) l V (Collett, 1902). 3 Scales in 1.1. 28 (Grieg, 1913)- Meristic characters. Radial formulae, vertebral and scale counts provided by various authors for the two species are shown in Table III. Although Collett origin- ally ascribed 6 elements to the first dorsal fin of both species, he later changed this to 212 P. J. MILLER 7 for G. orca. Lilljeborg (1884) and Smitt (1892) employed this supposed disparity in keys to Scandinavian gobies. L. orca is also regarded as having rather more articu- lated rays in the second dorsal and anal fins than L. scorpioides, and slight differences are indicated in pectoral and caudal fin ray counts. The number of scales along the lateral midline is reputed to be greater in L. scorpioides than in L. orca. For the orca and scorpioides groups of the present material the meristic values in Table IV have been obtained. Methods of counting are described in the systematic section below. To minimise the effect of regional variation, results for British and Scandinavian collections have been kept separate. Mean values are given for the more numerous British material. TABLE IV. Meristic Characters of Coloration Groups : Number of Observations (Highest in Bold Type) Against Value. M Mean Value for British Material Origin of specimens Coloration group orca scorpioides orca scorpioides orca scorpioides orca scorpioides orca scor orca scorpioides orca scorpioides British Isles 5 6 26 34 First dorsal fin rays Scandinavia 6 7 7 I 6 Second dorsal fin articulated rays 6 7 8 9 10 ii M 8 9 10 ii i 18 7 9-23 531 i 2 4 21 6 8-85 2 4 i Anal fin articulated rays 5 6 7 8 9 M. 56789 i 19 6 7 -19 54 32 22 7 6-97 i 4i Pectoral fin rays 15 16 17 18 19 20 21 M 17 18 19 20 21 2 15 20 15 18-92 742 i 4 7 9 30 10 i 18-56 4 2 Caudal fin branched rays 8 9 10 ii M 8 9 10 ii 24 i 9 -04 5 2 31 i 8-97 2 Scales in lateral series 25 26 27 28 29 M 25 26 27 28 29 2 18 10 6 2 26-68 i 3 2 i i 17 17 6 i 26-74 i i 2 Vertebrae including urostyle (Manx specimens only) 26 27 28 29 M 2 13 i 27-94 i 3 17 27-76 TAXONOMY AND BIOLOGY OF THE GENUS LEBETUS 213 Except for pectoral ray and scale counts in the Scandinavian fishes, where the number of observations is small, the most common value for each meristic character is the same in the two coloration groups and mean values are only slightly different. The wider range and somewhat lower means for fin ray counts in scorpioides are due 17- < 15- 13- CD O O CB O OO O O O o o o o o o o O oq o o e o o O OCX OOO p 1 | - 1 | - 1 15 1 I - 1 I 20 1 I 1 - 1 1 1 25 30 I I I I I I 15 20 25 30 Standard Length (mm) FIG. 20. Relation between standard length and length of last two rays of first dorsal fin (V, VI) and first two rays of second dorsal fin (I, i) in L. orca. Symbols as in Fig. 19. 242 P. J. MILLER eight diverse genera of gobies. The group is characterized by several peculiarities. Among these are the large non-osseous area between the symplectic, quadrate, and preoperculum ; the arrangement of the branchiostegals on the ceratohyal ; the absence of the parietal bones ; and the presence of a splintlike radial above and below the hypurals in the caudal skeleton. In all these features, the genus Lebetus appears as a typical gobioid fish, but there has been some doubt about the position of Lebetus among the families of this suborder. Of these about six are now recognized (Koumans, 1953 ; Gosline, 1955 ; Norman, MS.). The Rhyacichthyidae, Taenioididae, Kraemeriidae, and Microdesmidae are highly aberrant gobioids and need not be further dealt with in this discussion. The two largest and most important families are the Eleotridae and Gobiidae. Skeletal differences between these have been summarized by Regan (1911), and the diagnostic value of certain features reviewed by Gosline (1955). Nevertheless, in referring gobies to one or other of these families, it has been customary to use the external form of the pelvic fins as the chief guide in the usual absence of osteological data. According to Regan (1911) and other authors, the Eleotridae are characterized by the pelvic fins being separate, and the Gobiidae, when these fins are developed, by their union to produce a simple cup-shaped disc. However, various stages in the union or separation of the pelvic fins have been recognized among gobioid genera and the importance of this feature as a criterion of family position has recently been questioned, especially by Bohlke & Robins (1960^, b) who have found eleotrid, gobiid, and intermediate types of pelvic fins in a single genus (Coryphopterus Gill). In such cases as these, determination of systematic position within the suborder requires investigation of the skeleton. When erecting the genus Lebetus, Winther (1877) emphasized that the anterior membrane of the pelvic disc was missing, although the two fins were joined along their entire length in the midline. At first accepting (1885) the new genus, Collett (1896) was later of the opinion that intermediate stages in reduction of the anterior mem- brane could be found between the fully developed condition in Gobius and complete loss as reported for Lebetus, and doubted the validity of separating the latter from Gobius. On the other hand, Duncker (1928) found that the two pelvic fins were completely separated in Lebetus but stressed the desirability for re-examination of these fins in better preserved material in order to decide whether Lebetus belonged to the Eleotridae or the Gobiidae. De Buen (19306) associated Lebetus with two Mediterranean genera, one of which had previously been regarded as an eleotrid because of its almost completely separated pelvic fins, and placed this " Lebetus " group in the subfamily Gobiinae of the Gobiidae. Koumans (1931) similarly accepted Lebetus as a gobiid genus. In the present work, it has been confirmed that, in the least damaged examples, the pelvic fins of Lebetus are completely united between their origins and the distal extremity of the fifth ray of each fin, but are entirely without trace of an anterior membrane to complete the disc. To settle the question of family position with greater certainty, it became obvious that examination of the skeleton was necessary, and the results of this study reveal that Lebetus has the characters of a typical member of the Gobiidae. These include a T-shaped head to the palatine, loss of the mesopterygoid from the palatopterygoid arch, and absence of a hyper- coracoid in the pectoral girdle, where the laminar radials are inserted on the cleithrum TAXONOMY AND BIOLOGY OF THE GENUS LEBETUS 243 and the lowest also related to the hypocoracoid. In the possession of a spatulate glossohyal, five branchiostegous rays, a reduced metapterygoid, and loss of the opisthotic, Lebetus also reaches a higher level of specialization than that described among the eleotrids (Regan, 1911 ; Takagi, 1950 ; Gosline, 1955). Within the Gobiidae, the affinities of Lebetus undoubtedly lie with the subfamily Gobiinae, since the genus exhibits none of the more extreme modifications of the Sicydinae, Periophthalminae, Apocrypteinae, Tridentigerinae, Gobiodontinae, or Benthophilinae (Koumans, 1953 ; Norman, MS.). In the absence of an anterior membrane to the pelvic disc, Lebetus resembles a number of small Indo-Pacific genera i.e. Herreogobius Koumans, Quisquilius Jordan & Everman, Fusigobius Whitley, Amblyogobius Bleeker, and Zonogobius Bleeker (Koumans, 1953 ; Gosline, 1959) . With head and anterior part of body naked, elongate rays in the first dorsal fin, 22-30 scales in lateral series, and radial formula of D 2 1/8-9, A I/7~8 (Koumans, I 953)> Zonogobius appears to be closest of these to Lebetus. Differences comprise the larger gill openings, and more laterally compressed head in Zonogobius. However, it is impossible to trace relationships between European and exotic genera, when the arrangement of the lateral line system in so many of the latter, including Zonogobius, remains to be adequately described and figured. As shown in the normal illustration of Zonogobius corallinus sp. nov. (Mozambique) by Smith (1959, Fig. 29), the dis- tribution of sensory papillae on cheek and postorbital regions differs from that of Lebetus in the complete longitudinal rows and more numerous papillae. Koumans (1931) associated Lebetus with the genus Coryphopterus Gill ; as defined by Koumans, the latter included European Pomatoschistus species together with the genotype, C. glaucofraenum Gill, from the tropical Western Atlantic. In a revision of Coryphop- terus sensu stricto, Bohlke & Robins (19606) provide details of the cephalic lateral line system and other characters which do not point to any close affinity with Lebetus, in spite of the fact that two of their new Western Atlantic species (C. alloides and C. dicrus) are without the anterior pelvic membrane. In the European fauna there are two Mediterranean genera of gobies lacking anterior membranes to the pelvic fins. These are Odondebuenia and Cabotia, both introduced by De Buen (19306). Since Cabotia is preoccupied among the Lepidoptera, De Buen (i5th June, 1940) suggested Fagea as a replacement but was preceded in this by Whitley (May, 1940) who introduced Cabotichthys. The genus Odondebuenia contains two species which were originally (1907) placed among the Eleotridae as Eleotris balearicus Pellegrin & Fage and E. pruvoti Fage. Odondebuenia is character- ized by pelvic fins united only at their bases, naked nape and throat, unusual gill rakers (Fage, 1907, figs. 3 and 9), modified scales at the origin of the caudal fin, and meristic characters of D 2 1/9-11, A I/8-io, and Sc. 1.1. 24-32. Both species of Odondebuenia show sexual dimorphism in length of first dorsal fin rays (Fage, 1918 ; De Buen, 19306). The second of these Mediterranean genera, Cabotichthys, has the pelvics joined together for about half their length, nape and throat fully scaled, scales of the caudal peduncle not modified, and fin ray and scale counts of D 2 1/14, A 1/13, and Sc. 1.1.50. The genus is founded on one species, C. schmidti (De Buen) known only from the single type specimen (De Buen, 19306). De Buen (1931) included Odondebuenia, Cabotichthys and Lebetus in his " Lebetus " group of genera, the principal 244 P. J- MILLER character of which is the absence of the anterior pelvic membrane. Similarities in habi- tat also link these fishes, all three being obtained on coarse grounds where calcareous algae are noticeable constituents of the bottom deposits, and none exceed 50 mm. in length. On the face of the diagnoses given above, Lebetus would appear to be fairly closely related to Odondebuenia except for the occurrence in the latter of modified caudal scales and gill rakers, and the greater separation of the pelvic fins, while Cabotichthys stands quite distant from both in several respects. The lateral line system in Odonde- buenia and Cabotichthys has fortunately been investigated by De Buen (19306, 1940) and that of Lebetus is described above. Comparison of these accounts for the three genera reveals the heterogenous nature of their grouping. The patterns of sensory papillae and extent of the cephalic canals in Cabotichthys point to an intimate connec- tion with the genus Gobius, as defined by De Buen (19300, 1931), and, apart from the reputed form of the pelvics, other characters of this fish do not warrant any generic separation from Gobius. My own experience with dredged and trawled examples of Lebetus and other gobies is that the pelvic membranes are very susceptible to damage, and the pelvics of Cabotichthys, as figured by De Buen (19306, fig. 7) are reminiscent of such a condition. It may well prove that C. schmidti is no more than a damaged specimen of a Gobius species. The state of the pelvics in the Odondebuenia species seems well established from a number of specimens. In the arrangement of sensory papillae, this genus differs markedly from Lebetus in the greater number of papillae, and their distribution in well marked transverse and longitudinal rows on the cheek approaches that in Gobius, which Odondebuenia also resembles in the possession of a similar cephalic canal system. The morphological attributes common to both Odondebuenia and Lebetus are not of a very highly specialized nature and do not outweigh the considerable disparity in patterns of sensory papillae. Modification of scales at the base of the caudal fin involves pronounced elongation of all the ctenii in 0. pruvoti (Fage, 1907, fig. 7) or merely the lateral ctenii in 0. balearica (Fage, 1907, fig. n ; De Buen, 19306, fig. 2). Although not seen in Lebetus, it is of interest to note the occurrence of this peculiarity in the new gobioid genus Various, recently described from the West Indies by Robins & Bohlke (1961) and also with separated pelvic fins. These authors commented on the existence of comparable scales in the Californian eleotrid Chriolepis Gilbert and the gobiid Garmannia Jordan, which is represented on both sides of Central America and in the Caribbean. It is not proposed at this point to deal further with the relationships of Odondebuenia, except to call attention to the need for an osteological investigation of this genus. Loss of the anterior membrane from the pelvic disc has probably occurred on a number of different evolutionary lines among the Gobiidae and, as in the case of Odondebuenia and Lebetus, need not be a sure indication of close phyletic association. In geographical distribution Lebetus appears to be confined to the eastern Atlantic boreal region and may well have evolved there. It is in this area, therefore, that the closest relatives of the genus may be sought. Eight other gobioid genera are recorded from the eastern North Atlantic (De Buen, 1931) and all of these have a pelvic disc complete with anterior membrane. The two pelagic forms Aphya Risso and Crystal- TAXONOMY AND BIOLOGY OF THE GENUS LEBETUS 245 logobius Gill may be excluded from consideration on the grounds of extreme specializa- tion. Among the demersal gobies, the genera Deltentostem Gill, Pomatoschistus Gill, Chaparrudo De Buen, and Gobius L. exhibit various features of anatomy and lateral line organization (see De Buen, 1930^, 1931) which do not suggest any near relation- ship with Lebetus. The remaining genera, Lesueurigobius Whitley and Buenia Iljin, deserve more attention. The first of these, represented by Lesueurigobius friesii friesii (Collett), has a number of characters in common with Lebetus. Thus both genera are entirely without cephalic canals and in disposition of sensory papillae show a posterior extension of row a behind the eye, papillae in the oculoscapular furrow (row u), interorbital papillae, a high posterior termination for row i, and reduc- tion of rows m, n, and o (Sanzo, 1911, pi. 9, figs. 4, 5, as Gobius macrolepis ; De Buen, 1923, figs. 21, 22). However, the great abundance of papillae in Lesueurigobius contrasts with their relative scarcity in Lebetus, where several rows (g, x, z, c 2 , b, d) found in Lesueurigobius are little if at all developed, and others (i, e, ot, c-cp] inter- rupted. Except for reduction in rows m, n, and o, the resemblances listed may be accounted for by independent action of a similar evolutionary process in the past history of each genus. This was the replacement of existing cephalic canals by rows of sensory papillae. Considerable anatomical differences in size, habit, squamation, radial formulae, and coloration (Holt & Byrne, 1903 ; Duncker, 1928) would also suggest similarities are due to convergence, and that any affinity is relatively distant. No closer relationship can be demonstrated with Lesueurigobius sanzoi (De Buen) or L. lesueuri (Risso) from the Mediterranean (De Buen, 1923). The only genus now remaining is Buenia, the Atlantic species of which is B. Jeffrey sii (Giinther), and it is with this form that Lebetus appears at present to be most closely connected. Lacking scales on the head and predorsal regions, and with meristic characters of D 2 1/8-9, A I/y-8, and Sc. 1.1.25-30 (Duncker, 1928 ; personal observation), B. Jeffrey sii displays a resemblance to Lebetus which is also evident in the arrangement of the lateral line system (Text-fig. 21). On the cheek absence of transverse rows of papillae, and the short row of large papillae with an intermediate papilla between this and row a, recall the conditions in Lebetus. The lateral preorbital rows c 1 and c 2 are identical in both genera. On the dorsum of the head there is a correspondence in the reduced number and the arrangement of the papillae. Lebetus differs from B. Jeffrey sii in the disappearance of cephalic canals and their replacement by papillae of rows a 1 , u l , i 1 , and the anterior part of n, together with a certain " condensation " of other rows (d 1 , preoperculo-mandibular, opercular, and other series) and loss of rows d and b. The pattern of sensory papillae in Lebetus is obviously more specialized than that in Buenia, but the features present in both denote a greater affinity between these two gobies than is exhibited between Lebetus and other genera. In the persistence of cephalic canals and various rows of papillae, Buenia displays more ostensibly primitive characters than Lebetus. The former in addition retains the anterior pelvic membrane, is not so small as Lebetus, and does not show pronounced sexual dimorphism, although this does occur in growth of the first dorsal fin rays and probably in coloration of this fin. In the male genitalia, the testes are long and there is no free sperm duct. B. Jeffrey sii may accordingly show greater resemblance to the common stock from which the two are derived. The Mediterranean 246 P. J. MILLER species B. reticulatus (C.&V.) (=B. affinis (Kolombatovic)) is more removed from Lebetus in having a greater number of papillae especially in rows b and d, and a somewhat higher scale count (more than 33), than B. jeffreysii (Sanzo, 1911, pi. 9, figs. 9, 10 ; De Buen, 19300). In Manx waters B. jeffreysii has an offshore distribution FIG. 21. Sensory papillae and orifices of cephalic canals in Buenia jeffreysii, male, 31 mm. standard length. (A) Lateral, (B) dorsal views of head. Abbreviations as in Fig. 7. Orifices of canals cross-hatched, with greek lettering as in Sanzo (1911). like Lebetus and has even been taken from the same localities (see also Le Danois, 1910), but differences in precise habitat are perhaps reflected in the different body form and coloration (Holt & Byrne, 1903, fig. 4). Some of the characters of Lebetus may be interpreted as adaptations to a mode of life involving intimate association with dead shells, nodules of calcareous algae, and stones. The small size of the fish is obviously related to this type of habitat, and the normal coloration may have cryptic value against a background of Lithothamnion. Te Winkel (1935) has discussed those anatomical features of the gobiid Mistichthys TAXONOMY AND BIOLOGY OF THE GENUS LEBETUS 247 luzonensis which may be correlated with the extremely small size of this species at maturity (standard length 10-0-11-0 mm.). Although Lebetus is diminutive, it is appreciably larger than Mistichthys and the only obvious comparable modification in this genus is the reduced size of the testis, to which may be related the form of the urogenital papilla as considered above. A more detailed examination of the viscera than was attempted in the present study would be needed to investigate this question. Reduction in number of sensory papillae has already been mentioned, and may be linked with small body size (Barlow, 19616). The significance of loss of the cephalic canals is unknown ; these are also lacking in Lesueurigobius, which attains 100 mm. in L. friesii, but are present in smaller forms such as Odondebuenia, Buenia, etc. Another feature of probable adaptive importance is the absence of the anterior pelvic membrane, since this is also missing in Odondebuenia which appears to have a similar habitat and which in addition exhibits almost complete separation of the pelvic fins. While the exact affinities of Odondebuenia await investigation, it seems likely that in view of the resemblances in arrangement of sensory papillae between this genus and Gobius the condition of the pelvics is derived from an original gobiid disc structure in response to environmental demands, and is not primitive as in the Eleotrids. The advantages conferred by the alteration of the pelvic disc in Lebetus and Odondebuenia are not obvious. The loss of the anterior membrane seems to have occurred inde- pendantly in the evolution of the two genera as the exploitation of similar ecological niches proceeded in two areas from different stocks. It must be stressed that the above phylogenetic conclusions may at present be regarded as tentative. Not only does the position of Odondebuenia remain doubtful, but the alignment of Lebetus with Buenia may well be affected by future work on the lateral line system of exotic gobies and faunistic exploration in little known areas, which together could disclose closer resemblances and intermediate forms between these and Lebetus. SUMMARY The teleostean genus Lebetus Winther 1877 (Percomorphi-Gobioidea) has been studied from new Irish Sea material dredged off the south of the Isle of Man, and from other examples, including types, in the collections of various British and Scandi- navian museums. A redefinition of the genus includes the disposition of sensory papillae ; there is a reduction in number of the latter, and cephalic canals and an anterior membrane to the united pelvic fins are lacking. Two species were previously recognized : orca Collett 1874 and scorpioides Collett 1874. Among the various criteria coloration, meristic characters, body proportions used in the past to distinguish these two species, it was found that only in coloration and development of the dorsal fins could the present Lebetus material be divided into two groups, termed orca and scorpioides. The former consisted entirely of males, nearly all maturing or mature, the latter of females and immature males. The conclusion is reached that there is only one valid species, by page priority Lebetus orca. A detailed account of the external anatomy and osteology of this species is provided. In distribution L. orca is restricted to the European Atlantic boreal region, and a full list of records is provided in an appendix. The species is known chiefly from 248 P. J. MILLER coarse grounds, and has a wide bathymetric range from about 2 to 375 m. Investiga- tion of gut contents, using a points system of assessment, shows that L. orca is exclusively predatory, feeding on small demersal animals which largely comprise Crustacea and polychaetes. A number of endoparasites are listed. The male repro- ductive organs are peculiar in the relatively small size of the testis and the long free sperm duct ; " seminal vesicles " are present. Sexual differences exist in the form of the urogenital papilla, which in mature males has an unusual terminal circlet of vascularized papillae. In Manx waters, the breeding season probably commences in March and may extend to August. 140 to 270 ripening oocytes have been counted in mature females. Sexual dimorphism is found in coloration, and size of dorsal and anal fins. Examination of scales indicates that duration of life may be at least two years, with sexual maturation at one or two years. Maximum total length recorded is 39-0 mm. Investigation of life history and growth in the Manx specimens was complicated by the high selectivity of the sampling gear employed. In a discussion of the systematic position of Lebetus, it is shown that the skeleton is typically gobiid, and the arrangement of sensory papillae together with other features suggest that the nearest relative of this genus among the European gobies is Buenia Jeffrey sii (Giinther). Previous grouping of Lebetus with two Mediterranean genera lacking an anterior pelvic membrane (Odondebuenia De Buen and Cabotichthys Whitley) is criticized. The validity of Cabotichthys is doubted, and the suggestion is made that loss of the anterior membrane in Lebetus and Odondebuenia occurred independently during their evolution and occupation of a similar habitat in different areas. ACKNOWLEDGEMENTS I wish to thank Messrs J. S. Colman and A. B. Bowers, Marine Biological Station, Port Erin, and Mr. A. C. Wheeler, British Museum (Natural History), for their helpful criticism of this paper. My thanks are also due to Mr. D. Eggleston, Dr. R. G. Hartnoll and Dr. M. C. Miller for collecting specimens of Lebetus while dredging off the Isle of Man. For arranging the loan of other material, I am most grateful to Mr. E. K. Barth, Zoologisk Museum, Oslo ; Dr. E. Bertelsen, Carlsbergfondets Dana- Ekspeditioner, Charlottenlund Slot ; Mr. J. Nielsen, Universitetets Zoologiske Museum, Copenhagen ; Mr. A. C. Wheeler, British Museum (Natural History) ; and Mr. J. F. Willgohs, Zoologisk Museum, Bergen. Professor H. Brattstrom, Biological Station, Espegrend ; Dr. H. O. Bull, Dove Marine Laboratory, Cullercoats; Mr. B. Christiansen, Zoologisk Avdeling, Tromso Museum ; Dr. R. S. Clarke, Royal Scottish Museum, Edinburgh ; Mr. S. Mathiasson, Naturhistoriska Museet, Goteborg; Dr. B. B. Rae, Marine Laboratory, Aberdeen ; Dr. B. Swedmark, Kristinebergs Zoologiska Station, Fiskebackskil ; and Dr. A. E. J. Went, Fisheries Division of Dept. of Lands, Dublin, have kindly provided information on distribution or material present in collections under their care. Messrs J. W. Coles and S. Prudhoe, British Museum (Natural History), identified nematodes and platyhelminthes encountered in this study, while Dr. Eve C. Southward, Plymouth Laboratory, and Drs. N. S. Jones and D. I. Williamson, Port Erin Marine Biological Station, identified polychaetes, molluscs and arthropods among the gut contents. I am also most grateful to Mr. C. A. TAXONOMY AND BIOLOGY OF THE GENUS LEBETUS 249 Grainge, Zoology Dept., Liverpool University, for preparation of sections, and to Messrs P. J. Evennett, Zoology Dept., Leeds University, and D. J. Slinn, Port Erin Marine Biological Station, for photographic assistance. Most of the above investiga- tion was carried out during the tenure of a D.S.I.R. Research Studentship at the Marine Biological Station, Port Erin, and has been completed in the Zoology Dept., Glasgow University. I am therefore indebted to Mr. J. S. Colman and Professor C. M. Yonge, F.R.S., for provision of research facilities at Port Erin and Glasgow respectively. REFERENCES AFLALO, F. G. 1904. British Salt-water Fishes. 328 pp. London. ANDERSSON, K. A. 1942. Fiskar och Fiske i Norden. 1, 540 pp. Stockholm. BAL, D. V. 19400. A Study of Fish Eggs and Larvae from Manx Waters. 126 pp. Ph.D. Thesis, Liverpool University. 19406. Some recent additions of fish eggs and larvae to the fauna of Port Erin. Rep. mar. biol. Sta. 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(3), 20 : 288-291. 1888. Report on the fishes obtained by Mr. J. Murray in deep water on the north west coast of Scotland, between April 1887 and March 1888. Proc. roy. Soc. Edinburgh, 15 : 205-220. HARTNOLL, R. G. 1961. Studies on the Biology of Manx Spider Crabs. 177 pp. Ph.D. Thesis, Liverpool University. TAXONOMY AND BIOLOGY OF THE GENUS LEBETUS 251 HOLT, E. W. & BYRNE, L. W. 1898. Notes on the reproduction of teleostean fishes in the south-western district. /. mar. biol. Ass. U.K., 5 : 333-340. 1903- The British and Irish gobies. Rep. Sea Inl. Fish. Ireland 1901, Part 2 : 37-66. 1910. Fourth report on the fishes of the Irish Atlantic slope. List of recorded species, with references. Sci. Inv. Fish. Ireland 1908, No. 5, 7 pp. HYNES, H. B. N. 1950. The food of fresh-water sticklebacks (Gasterosteus aculeatus and Pygo- steus pungitius), with a review of methods used in studies of the food of fishes. J. anim Ecol 19 : 36-58. ILJIN, B. S. 1930. Le systeme des Gobiides. Trab. Inst. 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On the occurrence of Gobius orca, Collett, within the Clyde Sea Area. Trans, nat. Hist. Soc. Glasgow, 8 : 74-76. 252 P. J- MILLER PELLEGRIN, J. & FAGE, L. 1907. Description d'un Eleotris Mediterranean nouveau. Bull. Soc. zool. France, 32 : 11-12. PETERSEN, C. G. J. 1917. On the development of our common gobies (Gobius) from the egg to the adult stages etc. Rep. Dan. biol. Sta., 24 : 3-16. 1919. Our gobies (Gobiidae). From the egg to the adult stages. Ibid., 26 : 47-66. PETIT, G. 1941. Sur la morphologic cranienne de deux poissons cavernicoles aveugles : Typhleotris madagascariensis Petit et Coecobarbus geertsi Blgr. C.R. Fac. Sci. Marseille (i), 1 : 36-40. PRASAD, R. R. 1959. Reproduction in Clevelandia ios (Jordan and Gilbert), with an account of the embryonic and larval development. Proc. Nat. Inst. Sci. India, 25, B : 12-30. REGAN, C. T. 1911. The osteology and classification of the gobioid fishes. Ann. Mag. nat. Hist. (8), 8 : 729-733. ROBINS, C. R. & BOHLKE, J. E. 1961. A new gobioid fish from the Antilles and comments on Ctenogobius fasciatus and C. curtisi. Copeia 1961, No. i : 46-50. RUSSELL, F. S. i926. The vertical distribution of marine macroplankton. II. The pelagic young of teleostean fishes in the daytime in the Plymouth area, with a note on the eggs of certain species. /. mar. biol. Ass. U.K., 14 : 101-159. 19266. The vertical distribution of marine macroplankton. III. Diurnal observations on the pelagic young of teleostean fishes in the Plymouth area. Ibid., 14 : 387-414. I93oa. The vertical distribution of marine macroplankton. IX. The distribution of the pelagic young of teleostean fishes in the daytime in the Plymouth area. Ibid., 16 : 639-676. 19306. The seasonal abundance and distribution of the pelagic young of teleostean fishes caught in the ring-trawl in offshore waters in the Plymouth area. Ibid., 16 : 707-722. 1935. The seasonal abundance and distribution of the pelagic young of teleostean fishes caught in the ring-trawl in offshore waters in the Plymouth area. Part II. Ibid., 20 : 147-179. 1936. The seasonal abundance of the pelagic young of teleostean fishes in the Plymouth area. Part III. The year 1935, with a note on the conditions as shown by the occurrence of plankton indicators. Ibid., 20 : 595-604. 1939. On the seasonal abundance of young fish. VI. The year 1938. Ibid., 23 : 381-386. 1940. On the seasonal abundance of young fish. VII. The year 1939, January to August. Ibid., 24 : 265-270. SAEMUNDSSON, B. 1908. Oversigt over Islands fiske med oplysninger om deres forekomst, vigtigste biologiske forhold og okonomiske betydning. Skr. Komm. Havunders., No. 5, 140 pp. 1926. Fiskarnir (Pisces Islandiae.) Islenzk dyr, 1, 291 pp. 1927. Synopsis of the fishes of Iceland. Rit. Vis. Isl. Reykjavik, 2 : 1-66. 1949. Marine Pisces. Zool. Iceland, 4, Part 72, 150 pp. SANZO, L. 1911. Distribuzione della papille cutanee (organi ciatiformi) e suo valore sistematico nei Gobi. Mitt. zool. Sta. Neapel, 20 : 249-328. SCHMIDT, J. 1904. Fiskeriundersogelser ved Island og Faeroerne i sommeren 1903. Skr. Komm. Havunders., No. i, 148 pp. 1912. Introduction. Rep. Dan. oceanogr. Exped. Medit. 1908-1910, 1 : 1-49. SCHNAKENBECK, W. 1928. Rassenuntcrsuchungen am Hering. Ber. Deutsch. Wiss. Komm. Meeresf. (N.F.), 3 : 91-204. SMITH, J. E. 1932. The shell gravel deposits, and the infauna of the Eddystone grounds. /. mar. biol. Ass. U.K., 18 : 243-278. SMITH, J. L. B. 1951. The genus Paragobioides Kendall & Goldsborough 1911. Ann. Mag. nat. Hist. (12), 4 : 518-527. 1959- Gobioid fishes of the families Gobiidae, Periophthalmidae, Trypauchenidae, Taen- ioididae, and Kraemeriidae of the Western Indian Ocean. Ichthyol. Bull. Rhodes Univ., No. 13 : 185-225. SMITT, F. A. 1892. A History of Scandinavian Fishes. Parti. 566 pp. Stockholm. 1900. Preliminary notes on the arrangement of the genus Gobius, with an enumeration of its European species. Ofr. Svensk. Vet. Akad. Fork. 1899 : 543-555. TAXONOMY AND BIOLOGY OF THE GENUS LEBETUS 253 SOUTHWARD, E. C. 1957- The distribution of polychaeta in offshore deposits in the Irish Sea. J. mar. biol. Ass. U.K., 36 : 49-75. SPRINGER, V. G. & MCERLEAN, A. J. 1961. Spawning seasons and growth of the code goby, Gobiosoma robustum (Pisces : Gobiidae), in the Tampa Bay area. Tulane Stud. Zool., 9 : 87-98. TAKAGI, K. 1950. On the glossohyal bone of the gobioid fishes of Japan, with some phylo- genetic considerations. Jap. J. Ichthyol., 1 : 37-52. TANING, A. V. 1940- Island-Faera-Omraadets Kutlinger (Gobiidae). Vid. Medd. Dansk. naturh. Foren., 104 : 267-282. TE WINKEL, L. E. 1935- A study of Mistichthys luzonensis with special reference to conditions correlated with reduced size. /. Morph., 58 : 463-535. VIVIEN, J. H. 1939- fitude preliminaire du tractus genital chez Gobius paganellus L. Trav. Sta. zool. Wimereux, 13 : 713-721. WENT, A. E. J. 1957. List of Irish Fishes. 31 pp. Dublin. WHITLEY, G. P. 1930. Additions to the check-list of the fishes of New South Wales (No. 3). Austr. Zool. Sydney, 6 : 117-123. - 1931. Studies in ichthyology, No. 5. Rec. Austr. Mus. Sydney, 18 : 138-160. 1940. The Nomenclator Zoologicus and some new fish names. Austr. Nat. Sydney 1940 : 241-243. WINTHER, G. 1877. Om de danske fiske af slaegten Gobius. Naturh. Tidsskr. Kjobenhavn (3), 11 : 41-56. 1879. Prodromus ichthyologiae danicae marinae : fortegnelse over de i de danske farvande hidtil fundne fiske. Ibid., 12 : 1-68. APPENDIX Distribution of Lebetus orca A list of published and unpublished records known to the author is given below. A complete sequence of information is as follows : locality, number of specimens, sex, museum number, standard length + caudal fin length or total length in millimetres (mm.), depth of water in metres (m.), nature of ground, date of capture, and authority in the case of published records. When a number of records from the same locality or area are given in the one publication, these are separated by semicolons and the authority placed at the end. Abbreviations employed are : c., coarse ground ; cl., clay ; d., damaged; gr., gravel; h., hard ground ; Lith., Lithothamnion ; m., mud ; midw., midwater ; Mod., Modiolus ; n.d., no date ; s., sand ; sh., shells ; St., stones ; surf., surface ; t.L, total length ; w., weed. ICELAND : Off S. and W. coasts, Eystra Horn to BreiSifjordur, " pelagic fry . . . rather rarely " (Taning, 1940; Saemundsson, 1949). FAEROES : Off N. coast, and Faero Bank, postlarvae (Taning, 1940). ROCKALL : On and near Rockall Bank, 1908 (Tailing, 1940). NORWAY Hemnefjord (Trondelag) : one