Bijdragen tot de Dierkunde, 56 (2): 214-220 — 1986
Notes on the distribution and phylogenetic significance
of post-cloacal sacs and bones as occurring in
the Gekkota (Reptilia)
by
J. Bastinck
do Institute of Taxonomic Zoology, P.O. Box 20125, 1000 HC Amsterdam, The Netherlands
this which Abstract cloacal sac variability in genus, was
first noted by Hoogmoed (1973), is here con- On the basis of Kluge’s (1982) paper on post-cloacal sacs firmed and commented upon. and bones in the Gekkota the following matters are
the situation in the use of the names Gekkota and Gek- presented: (1) gekkonid genera Kluge's Microscalabotes and Paragehyra is recorded, (2) the situation konoidea, and his interpretation of the in the is reviewed gekkonid genus Lygodactylus extensively of phylogenetic significance post-cloacal sacs, and commented the upon, (3) post-cloacal sac variability here commented well. are upon as in the first noted gekkonid genus Thecadactylus, by
Hoogmoed (1973) and evidently overlooked by Kluge, is confirmed and commented upon, (4) Kluge’s interpreta- RESULTS AND DISCUSSION tion of the phylogenetic significance of post-cloacal sacs is discussed. USE OF THE NAMES GEKKOTA AND GEKKONOIDEA
used the name Gekkota sensu Zusammenfassung Kluge (1967: 10)
Underwood (1957). According to Underwood Anhand Kluges (1982) Veröffentlichung über Post- the infra-order Gekkota consists of the families kloakalspalten und -knochen bei den Gekkota, wird Eublepharidae, Gekkonidae, Sphaerodac- Folgendes behandelt: (1) die Situation innerhalb der three families Gekkoniden-Gattungen Microscalabotes und Paragehyra wird tylidae (these collectively dokumentiert, (2) die Situation innerhalb der Gekko- Gekkonidae regarded as one family by Kluge) wird kritisch niden-Gattung Lygodactylus zusam- take it that and Pygopodidae. We may then, mengefasst, (3) die Variabilität der Postkloakalspalten bei whenever Kluge is using the name Gekkota he der Gekkoniden-Gattung Thecadactylus, erstmalig is referring to the Gekkonidae and Pygopodidae wahrgenommen von Hoogmoed (1973) und offenbar
übersehen wird und von Kluge, bestätigt besprochen, (4) collectively.
Kluges Interpretation der phylogenetischen Bedeutung also used the Kluge name Gekkonoidea. von Postkloakalspalten wird diskutiert. Perusal of his paper shows that Kluge is not
referring here to the Gekkonoideasensu Under- INTRODUCTION in of wood, as could have been expected view
review his but Kluge (1982) published an extensive of use of the name Gekkota, to the Gek-
of and bones in konoidea the occurrence post-cloacal sacs sensu Hoffstetter (1964) (see Kluge, the Gekkota. In that paper Kluge noted that he 1967: 12). Hoffstetter considers the superfami- had been able the Gekkonoidea contain the families not to study gekkonid genera ly to
Microscalabotes and Paragehyra, and that the situa- Ardeosauridae (|) and Gekkonidae (and, ten- tion in the gekkonid genus Lygodactylus required tatively, Bavarisauridae (f) and Palaeolacer-
I able tidae further study. was to study these three (f)). This would mean that Kluge, by
and I to some extent observa- the name is refer- genera put my using Gekkonoidea, simply tions record here. Gekkonidae far families on ring to the as as recent
A concerned. remarkable omission in Kluge's otherwise are One gets the strong impression,
work is formed the that considers the Gekkonoidea very comprehensive by however, Kluge
situation in of the and peculiar regarding post-cloacal sacs to consist Gekkonidae Pygopodidae.
the The is of gekkonid genus Thecadactylus. post- This notion fully supported by Kluge's use
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the name Gekkonoidea in later works (Kluge, In order to investigate the presence/absence
1976; 1982). Reference to Underwood shows of post-cloacal sacs and bones in this species,
that this author considers the Gekkonoidea to Hoogmoed (1973) examined 39 specimens. He
contain the Eublepharidae, Gekkonidae and found 21 specimens (o*C + 9 9) in which sacs
Sphaerodactylidae, which means that Kluge's were present and 17 specimens (o*cc + 9 9) in
of is which these absent use the name Gekkonoidea evidently not structures were (in one
Underwood's definition For he based on either. juvenile specimen could not decide whether
this novel Gek- He all males apparently use of the name sacs were present or not). found to
konoidea fails line of Kluge to give any reason- be in possession of post-cloacal bones.
cite of the I examined 17 of T. ing or to any authority. Kluge's use specimens rapicauda
names Gekkota and Gekkonoidea means that in them 5 of the myself, among specimens already
effect from he is treating them as synonyms. examined by Hoogmoed. Apart 1
In the remainder of this the Gek- in which I could paper name specimen not unequivocally
used determine whether I kota will be sensu Underwood; the name sacs were present or not,
Gekkonoidea will not be used. found 8 specimens (4 CO", 4 9 9) in which sacs
and 8 3 in were present specimens (5 CO", 9 9)
which absent. All in DISTRIBUTION OF POST-CLOACAL SACS AND BONES they were males were
IN THE GEKKONIDAE of I thus possession post-cloacal bones. can con-
firm Hoogmoed's observations. Introduction The above-mentionedobservations show that
who examined According to Kluge (1982), the Gekkoninae there is least among at one almost all constituting the family Gek- genera genus in which post-cloacal sacs are not konidae, post-cloacal sacs and bones are invariably present or absent; in fact, this situa- distributed as follows: tion be the Gek- appears to unique among (1) in all of the Diplodactylinae present genera konidae in general. Apparently, Kluge and Eublepharinae; overlooked this variability in Thecadactylus absent of in all genera the Sphaerodac- and/or of (2) was not aware Hoogmoed's obser-
tylinae; vations.
in the (3) variably present Gekkoninae.
Kluge listed the following taxa belonging to
the Gekkoninae characterized the The situation in Microscalabotes and as being by
absence of both and bones: the post-cloacal sacs Paragehyra
genera Aristelliger, Asaccus, Millotisaurus, As noted above, Kluge (1982) stated that he did Narudasia, Pristurus, Quedenfeldtia and Saurodac- examine the Microscalabotes and not genera and of the tylus, two species genus Phyllodactylus, Paragehyra in the course of his survey of the Gek- viz., P. riebeckii and P. (Peters, 1882), koninae. trachyrhinus 1899. Boulenger, The sole of the Microscalabotes species genus Kluge noted that the situation in Lygodactylus Boulenger, 1883 is M. bivittis (Peters, 1883), required further study and declared not to have which is known from a very few specimens examined the genera Microscalabotes and (Pasteur, 1967). According to Pasteur (1964b) Paragehyra. the is characterized the absence of genus by
post-cloacal sacs; Pasteur does not provide The situation in Thecadactylus information concerning post-cloacal bones. I
One of the listed able M. genera by Kluge (1982) as was to study the single specimen of
and bivittis that is in the collection of the possessing post-cloacal sacs (0'0' + 9 9) present
bones is 1817. This Paris (CCf) Thecadactylus Oken, museum (MNHNP A.61). Although the
of T. is in bad it is clear that genus is composed a single species: specimen a rather state,
I thus rapicauda (Houttuyn, 1782). no post-cloacal sacs are present; can con-
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firm Pasteur's I could (1964b) statement. not genus is apparently very close to Agamura (De
detect the of bones. The I that it presence post-cloacal Witte, 1980), suspect Kluge treats as a
absence of bones is corroborated the of the latter. It should be by junior synonym noted
of series of that the has presence a well-developed preanal up to now genus Rhinogekko not
it that the been pores, which makes extremely likely formally suppressed.
in which Szczerbak specimen is a male, case post-cloacal Golubev & (1981) erected the
if should have been detec- accommodate the bones, present, easily genus Carinatogecko to species
these known table (in the bone-bearing genera struc- originally as Bunopus aspratilis Anderson, tures are invariably present in males). 1973 and Tropiocolotes heteropholis Minton,
is another Apparently, Microscalabotes member Anderson & Anderson, 1970. It seems possible
the series of that of the of gekkonid genera characterized Kluge was unaware description of
the absence of and bones. this the time he by post-cloacal sacs genus at was writing his 1982
The genus Paragehyra Angel, 1929 likewise is paper. However, since both Bunopus and
its P. monotypic; only member, petiti Angel, Tropiocolotes were found by Kluge to be in
1929, is known only from the holotype. This possession of both post-cloacal sacs and bones, specimen was studied by Brongersma (1934), and since C. aspratilis and C. heteropholis do not who observed that post-cloacal sacs were pres- seem to differ from their former respective con-
bones not. in in this ent, while post-cloacal were I was geners respect (Anderson, 1973; Minton the of P. it certain that position to study the holotype petiti et al., 1970), seems Carinatogecko
Post-cloacal is characterized the of these (MNHNP 29-75) myself. sacs are too by presence present in this specimen indeed, albeit rather structures. weakly developed. I could not detect the The situation in Lygodactylus will be discussed of bones. presence post-cloacal Thus, the at end of this paper. observations Brongersma's apparently were correct. The evident absence of bones, the weak DISTRIBUTION OF POST-CLOACAL SACS AND BONES development of the sacs (in males sacs are IN THE PYGOPODIDAE almost always strongly developed), and the lack examined all of the Kluge (1982) genera of a swollen tail-base make it noticeably very Pygopodidae and found post-cloacal bones to be likely that the specimen is a female. It is to be in all this that when present genera constituting family. expected males, found, will prove to Post-cloacal sacs were found to be be in of both and variably possession post-cloacal sacs present in the This variability is bones. It therefore that far Pygopodidae. appears as as post- summarized in table I. cloacal and bones concerned the sacs are genus
Paragehyra does not differ from the majority of Table I gekkonid genera. Distribution of the of the post-cloacal sacs among genera
Pygopodidae according to Kluge (1982).
Genera not mentioned by Kluge o*o* 99
Two genera belonging to the Gekkoninae were Aclys ? not mentioned either as by Kluge (1982) variable Aprasia present studied or not studied; these are Rhinogekko and Delma absent absent
Carinatogecko. Lialis absent absent
The Ophidiocephalus variable variable genus Rhinogekko was erected by De Paradelma in 1973 and present present Witte contains one species: R. Pletholax absent present (?) misonnei De 1973. the Witte, Up to now genus Pygopus present present has only been mentioned in literature by De
Witte himself from a non- (1973, 1980), apart 99 Aprasia display inter- and intraspecific variation; reference in Welch As informative (1983). this CTCf + 99 Ophidiocephalus display intraspecific variation.
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SIGNIFICANCE OF Even and bones PHYLOGENETIC POST-CLOACAL if it were supposed that sacs
BONES have been lost only once in the phylogeny of the
Gekkonidae, it would be difficult to regard the From his in observations on post-cloacal bones combined loss of such closely associated struc- the Gekkonidae, Pygopodidae, Xantusiidae tures as coincidental. It seems certain, how- and the extinct protorosaurians Tanystropheus in and bones ever, that the Gekkonidae sacs and Tanytrachelos, Kluge (1982) draws the have been lost several times: Russell (1977) following conclusions, with which I completely that loss of and convincingly suggests sacs agree: three bones may have occurred at least times, (1) post-cloacal bones in the Gekkonidae and while maintains that loss Kluge (1982) may Pygopodidae are homologous structures; have occurred from three to six times (2) post-cloacal bones in the Gekkota are not independently! Under the afore-mentioned homologous with either those in the Xan- that hypothesis we are asked to believe these tusiidae or those in Tanystropheus and Tany- combined losses are due solely to coincidence. trachelos; This is asking too much of anyone's credulity. (3) post-cloacal bones in the Gekkota are a An alternative hypothesis suggests itself when unique synapomorphy. considering the following facts:
If bones in 1. post-cloacal are present a given PHYLOGENETIC SIGNIFICANCE OF POST-CLOACAL sacs may be SACS species, post-cloacal this (a) invariably present in same species;
In the suborder Sauria post-cloacal sacs are a e.g. Hemidactylus sp.;
the in unique feature of Gekkota: they occur only (b) variably present this same species;
in the Gekkonidae and Pygopodidae (Kluge, e.g. Thecadactylus rapicauda;
of his observations in this 1967). On the basis Kluge (c) invariably absent same species;
draws the with Pristurus (1982) following conclusions, e.g. sp.
which I 2. If bones are absent in a fully agree: post-cloacal given
in the Gekkonidae and absent (1) post-cloacal sacs species, post-cloacal sacs are too; e.g.
Pristurus Pygopodidae are homologous structures; sp.
(2) post-cloacal sacs in the Gekkota are a These observations lead to the conclusion that
within the of is unique synapomorphy. a given species presence bones
In view of the in for the of Once post-cloacal sac variability a prerequisite presence sacs.
the stated that the of Pygopodidae (see table I) Kluge bones are present, presence/absence sacs
"fluctuate". "the fact that cloacal bones and sacs do not may Once bones are absent, sacs
is sufficient of absent This always co-occur in pygopodids are necessity too. interpretation
for them char- fit the known facts better than justification treating as separate seems to Kluge's
acters in the Gekkonoidea" (Kluge, 1982: 349). hypothesis.
In the of the other words, Kluge here suggests that post- Thus, following picture phylogeny
and of cloacal sacs bones (as occurring in the Gek- gekkotan post-cloacal sacs and bones
The kota) are phylogenetically non-correlated char- emerges. ancestral gekkotan was in posses-
when the and Within acters. This hypothesis gains strength sion of both sacs bones. the
in the (intraspecific) post-cloacal sac variability Pygopodidae bones were consistently retained, gekkonid genus Thecadactylus (which Kluge while the presence of sacs became variable (and
in overlooked) is also taken into account. even nonexistent) a number of genera.
The of the above-mentioned Within the Gekkonidae consequences post-cloacal sac
difficult hypothesis, however, are to accept. variability evolved only once (in Thecadactylus),
Under Kluge's hypothesis the combined loss of while loss of both bones and the closely both and bones in 15 associated occurred least but post-cloacal sacs gekkonid sacs at once, prob-
coincidental. from three six times. genera must be regarded as purely ably to
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THE IN cloacal SITUATION LYGODACTYLUS sacs. All males were in possession of
easily detectable post-cloacal bones.
Pasteur divided the it should be clear that the (1964a) genus Lygodactylus Thus, now genus
Gray, 1864 into two subgenera: Lygodactylus, Lygodactylus exhibits intrageneric (but not
characterized by the absence of post-cloacal intraspecific) variation regarding the
and characterized the of sacs, Domerguella, by presence/absence post-cloacal sacs and
of bones. This this presence post-cloacal sacs. Up to now, means that type of intrageneric
Pasteur has not supplied any information con- variation is now known from two gekkonid
the of and cerning presence/absence post-cloacal genera (Lygodactylus Phyllodactylus (see
bones in the The genus Lygodactylus. subgenus above)).
contains some the The variation within the Lygodactylus forty species; genus Lygodactylus
subgenus Domerguella consists of five, viz., L. might be indicative of generic status of the
Pasteur L. expectatus & Blanc, 1967, guibei subgenera Domerguella and Lygodactylus, since
L. and bone Pasteur, 1964, madagascariensis (Boettger, intrageneric post-cloacal sac
L. and L. is in the Gek- 1881), miops Gunther, 1891, rarus variability evidently very rare
Pasteur & 1973. The konidae. Blanc, subgenus Osteological comparisons might sup-
Domerguella is strictly Madagascan in ply the information needed for the justification
distribution. of least this method recognizing two genera (at
stated that the excellent results in the of the Kluge (1982) genus Lygodac- produced case
is characterized in its the erection of the whose tylus entirety by genus Asaccus, species
absence of post-cloacal sacs and bones. How- were formerly assigned to Phyllodactylus (Dixon
he also stated that the situation in the ever, & Anderson, 1973)).
further genus Lygodactylus required study. According to Pasteur (1964b) the presence of
Unfortunately, Kluge supplied no list of the post-cloacal sacs (and, as mentioned above,
species he examined; in earlier work (Kluge, bones) in the subgenus Domerguella is to be
of 1967) he mentioned L. capensis (Smith, 1849), regarded as a case reappearance of these
in L. conraui Tornier, 1902, and L. picturatus structures, other words, as a case of evolu-
reversal. (Peters, 1870) as studied. tionary
In order of the the to clarify some confusion con- Generally, occurrence of evolutionary
the situation in the reversal is not cerning genus Lygodactylus, regarded as a common
I examined representatives of the following phenomenon (Dollo's law), but as far as the
the the species belonging to subgenus Lygodactylus: situation in genus Lygodactylus is concerned,
L. L. L. Pasteur certain his In the capensis, conraui, fischeri Boulenger, seems to be of case.
1890, L. luteopicturatus Pasteur, 1964, L. ocellatus above-mentioned publication he presented, in
Roux, 1907 (including the lectotype, ZMA addition to the occurrence of post-cloacal sacs
11347), L. picturatus, L. robustus Boettger, 1913, (and bones), a series of eight characteristics of
L. tuberifer Boettger, 1913, and L. verticillatus the subgenus Domerguella that should be
1895. Neither indicative of advanced Mocquard, post-cloacal sacs nor regarded as a very state bones could be detected in of the of this the any specimens subgenus (as compared to subgenus examined. Lygodactylus). Furthermore, during a recent
I also conversation this assured examined representatives of the five on matter Pasteur me species of the subgenus Domerguella: 2 crcr + 2 that he was "absolutely convinced" that the
of L. the BP of sacs and bones in the 9 9 expectatus (including holotype, presence subgenus
1 of L. of L. be of 640), C + 2 9 9 guibei, 1 O" + 2 9 9 Domerguella should regarded as the result madagascariensis, 2 crcr of L. miops and 1 9 (the evolutionary reversal.
BP of All Two for the in holotype, 1.72) L. rarus. specimens possible explanations situation
in of discernible the without were possession clearly post- genusLygodactylus, necessitating use
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REFERENCES of the concept of evolutionary reversal, suggest
themselves: ANDERSON, S. C., 1973. A new species of Bunopus (Rep-
the the Iran and a lizards of (1) within genus, subgenus Domerguella tilia: Gekkonidae) from key to
is the Bunopus. 29 (4): 355-358. the more primitive one (subgenus genus Herpetologica, BRONGERSMA, L. D., 1934. Contributions to Indo- Lygodactylus advanced); Australian herpetology. Zool. Meded. Leiden, 17 (3- (2) the situation within the genus is an example 4): 161-251, pis. I-II. (Also published separately as of the of mosaic evolution: the phenomenon Ph.D. thesis, University of Leiden, by E. J. Brill,
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DIXON, R. & S. C. ANDERSON, 1973. A new and diverged from a common ancestor which J. genus species of gecko (Sauria: Gekkonidae) from Iran and was in the possession of both post-cloacal Iraq. Bull, south. Calif. Acad. Sci., 72 (3): 155-160. sacs and bones. In the subgenus N. N. 1981. Carinato- GOLUBEV, M. L. & SZCZERBAK,
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Russian.) were retained, while in the otherwise less HOFFSTETTER, R., 1964. Les Sauria du Jurassique divergent subgenus Lygodactylus they were superieur et specialement les Gekkota de Baviere et lost. de Mandchourie. Senckenberg. biol., 45 (3/5): Both in direct contradiction explanations are 281-324.
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This is , 1976. A reinvestigation of the abdominal not the place to enter into an extended
musculature of gekkonoid lizards and its bearing on discussion of Pasteur's detailed and carefully their phylogenetic relationships. Herpetologica, 32 compiled work. It should here be noted that a (3): 295-298. of comparative histological investigation gek- 1982. Cloacal bones and sacs as evidence of , gek- konid and bones shed post-cloacal sacs might konoid lizard relationships. Herpetologica, 38 (3):
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Received: 21 February 1986
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