Of the 12 species groups, all the species within each of five groups containing from three to nine species are allopatric. One of the five species in one group is broadly sympatric with six of the eight species in the other group. Such distribution patterns appear to be better explained by the tectonic history of the relevant areas than by dispersal.
Distribution of the Oculus-Yaeyamaensis Clade 122 Appendix II: Relationships and Species Biogeography of. Thus, the distributions of these species would be less well known than those of the species of the other three genera. The apparent presence of a planktonic phase suggests that species of Ecsenius should, in general, have distributions no less restricted than those of species of Entomacrodus, Omobranchus, and Istiblennius.
I believe that the explanations for many of these distribution patterns lie in the tectonic history of the Indo-Pacific, which is only now being unraveled. A missing dorsal fin spine is recognizable when a pterygiophore in the spiny portion of the fin does not support any element.
Most of the species of Ecsenius can be identified by color pattern alone, and I believe that relationships are largely reflected by color-pattern similarities. Often, color pattern characteristics are unclear in printed illustrations, and published colors may be very different from the original color photograph. The colors of the images obtained therefore do not appear in the photographs as they would to an underwater observer under natural conditions.
Because the phylogeny of the Salariini is unresolved, I am unable to unambiguously polarize the characters for Ecsenius species. Furthermore, many of the characters I had for consideration, especially at the species level, are modal states for meristic characters, which I find impossible to unambiguously polarize (one exception: the number of segmented caudal fin rays). In several of the groups established early in the study, all species were allopatric.
Most of the halftone drawings, and some halftone photographs, of fish have been published and acknowledged in my previous studies, but all are listed separately in brackets in the captions of the illustrations. Springer (1968) implicitly assumed the monophyly of the Blenniidae, and I reaffirm that the three main characters used to define the family are synapomorphies.
VERTEBRA
Randall (BPBM) who allowed me to use a very large number of his original color photographs, and G.R. Publications, Inc., who generously provided the color separation and printed and donated the color plates for inclusion in the final publication. Portions of early drafts of this manuscript were discussed or read by several persons, notably R.H.
Horn, for the great effort they put into improving format and text and bringing the whole thing into a printable state. Important new material, added since McKinney and Springer (1976), resulted from my fieldwork in Fiji, Philippines, Rotuma, Papua New Guinea, Pohnpei, and Cargados Carajos. My fieldwork in these places was supported by grants from the Dreyfus Foundation and the Lewis L. and Rosa Strauss Memorial Foundation.
LIGAMENT
EPIPLEURAL RIB
Springer and Smith-Vaniz also implicitly hypothesized a third synapomorphy of Ecsenius: the presence of posterior and anterior canine teeth (Figure 2). While the combined presence of both types of canines is unique to Ecsenius among the Blenniidae, each type of canine must be treated as a separate character. Superficially, the presence of anterior canines in Ecsenius seems to indicate that Ecsenius is the sister group of all other Salariines and that it is plesiomorphic for anterior canines.
However, the polarity of the anterior canines in Ecsenius is ambiguous, because the Parablenniini exhibit both conditions for the character. Except for Ecsenius midas, which has well-differentiated anterior canines, the anterior canines in non-skeletonized preparations of Ecsenius species are either so modified that they superficially appear to be incisors, or they are barely recognizably differentiated from the incisors (Figure 2 illustrate) skeletal lower jaw of E. bicolor, in which canines are slightly differentiated;.
DENTARY
POSTERIOR CANINE
ANTERIOR CANINE
INCISOR REPLACEMENT INCISOR
The possibility that the presence of anterior canines in Ecsenius is a homoplasic condition, however, excludes the unequivocal hypothesis of a primitive condition of these canines in Ecsenius. Posterior canine teeth are present in Blenniidae in only 17 of the 21 genera of Salariini, but may not be present in all species of the genus or in both sexes of the species. However, I believe that dogs are a synapomorphy of the Salariinae, which have been secondarily lost in four otherwise highly specialized genera consisting of five species that do not have them.
Two synapomorphies of Ecsenia—loss of lateral extrascapulars (or their fusion with pterotics, Figs 3 and 4) and filiform caudal fin lobes—occur differentially in some genera of the tribes Omobranchini and Nemophini, which are not sister groups to the Salariini. (Smith-Vaniz, 1976). GENERAL BIOGEOGRAPHIC COMMENTS.—The distribution of many species and groups of species of Ecsenius is closely demarcated (Figs. 5-13). This circumstance exists even though the geographically closer species may not be the sister group of the species in question.
A general breakdown of the distribution of the species and species groups is given in Table 5. The base maps for all but one of these figures are slightly modified from those used by Springer (1982), which include dashed lines indicating generalized margins of the major lithospheric plates.
Q ECSENIUS AEQUALIS Q ECSENIUS MANDIBULARIS
0 ECSENIUS KURTI O ECSENIUS SCHROEDERI
Q ECSENIUS LUBBOCKI Q ECSENIUS TRILINEATUS
51.0 females 1
There are gaps, from which no species of the Oculus group are known, between the distributions of E. I believe that the nape spots are either a synapomorphy of the Oculus group or homplastic in E. The Batan Islands are politically part of the Philippines, but based on the presence of E.
The most prominent part of the pattern appears on the ventral surface of the head. In the other three species of the Yaeyamaensis group, most individuals have a dark band at the base of the pectoral fin. Individuals of all species of the Yaeyamaensis group usually have a visible dark spot on the inner (axillary) surface of the fleshy base of the pectoral fin.
Another dark spot dorsal to the axilla of the pectoral fin is also probably a remnant of the postorbital stripe. Pale spots on the body are less noticeable than those of the other species of the Yaeyamaensis group. The cheeks below the level of the midpostorbital stripe are never punctured with fine, dark spots.
In this specimen, the dark pigments in the head and body are shades of brown (mid-postorbital and pectoral-fin basal stripes are almost black). The dark opercular line is dark and distinct only on the pale ventral surface of the head.
Komodo 1 3
There is a dark spot on the axillary surface (inside) of the fleshy pectoral fin base in E. A discrete axillary spot is otherwise only found in the species of the Yaeyamaensis Group and E. Cohen (1973), in an admittedly limited analysis. of endemism among coastal fish in the Indo-Pacific, noted that a relatively high percentage (22%) of fish in the western Indian Ocean are endemic.
