Proposal
(15) to South American
Classification Committee
Elevate Butorides
sundevalli (Lava Heron) to species rank
Effect on South American
CL: this proposal would split the Striated Heron, Butorides
striatus, into two species, B. striatus and B. sundevalli.
Background: Two
forms of Butorides herons breed in the Galapagos Islands, a typical pale
taxon evidently indistinguishable from mainland B. striatus, and an
endemic dark taxon, sundevalli. These are either treated as conspecific
(e.g., Martínez-Vilata and Motis
1992), as is the current status in our baseline list, or as separate species
(Harris 1982). The endemic taxon is a common nester on all islands (Harris
1973, 1982); striatus-like birds are known only from seven islands.
Here's the direct quote
from Harris (1973):
"The
situation regarding these species is confused. B. sundevalli is
endemic and common in coastal areas of all the islands, and until recently
there had been no suggestion that another Butorides sp. ever
occurred in the Galápagos. However, birds resembling B. striatus have
been recorded on Isabela (nesting), Santa Cruz (nesting), Fernandina, Duncan,
Santa Cruz, San Cristobál, and Pinta. Present
knowledge is insufficient to ascertain whether there are two species of Butorides breeding
in Galápagos which hybridize (as intermediates occur) or whether there is one,
very variable species. If B. striatus does indeed breed, it is
not a recent colonizer since skins assignable to the species were collected by
the Academy Expedition 1905-6 (L. C. Binford, unpubl. data).”
Payne (1974) examined 58
specimens from the Galapagos and concluded the following:
" show all possible intermediate
plumage colours." " high proportion of
intermediate birds indicates considerable successful breeding and genetic
recombination." " The Galapagos herons are variable, but nearly all
adults are separable from adults of continental populations." " The
Lava Heron is probably best considered a distinct subspecies, B.
striatus sundevalli, at least until studies on breeding pairs
have been made on the islands."
Payne's table indicates
that only 13 are really "pure" Lava Heron phenotype, and 18 have dark
gray or gray underparts, darker than South American striatus but paler
than typical "Lava Heron." Only two were as pale as South
American striatus, but can still be distinguished from them. Payne
speculated that the variability might be the result of multiple invasions of
the Galapagos by striatus stock at different times.
Kushlan (1983), who
studied the breeding behavior of Lava Herons (but evidently not the local
Striated-like phenotype), noted a number of subtle differences in behavior
between Lava Herons and what had been published for South American B.
striatus but concluded:
"Pair formation of the Lava Heron generally resembled
that of the Green-backed Heron described by Meyerriecks (1960). The substantial
similarities are added evidence of a close relationship and lend no support for
doubting Payne's (1974) view of the conspecificity of the three forms on Butorides."
Kushlan noted the
following differences in sundevalli from striatus: (1) female
performs stretch at nest site, (2) does bill snaps, and (3) doesn't sway.
Additional minor differences were also noted. However, he emphasized that his N
was small on Galapagos populations studies, and that unstudied populations of
year-round resident striatus populations there or elsewhere might also
show these differences (which he interpreted as having to do with year-round
territoriality and nesting three times per year).
Kushlan also noted
differences between the breeding bare part colors of the two, but seemed not to
attach significance to them. Lava legs turn from gray to reddish-orange,
whereas Striated (and Green) legs go from orange to orange-red. Lava lores go
from green to bright cobalt blue, whereas in Striated, they turn blue-black.
[For those without access to color plates, note that sundevalli is
almost all black, by far the most distinct plumage phenotype in the
global striatus group.]
Recommendation: I will
vote NO on my own proposal, which began as an investigation that I thought
would conclude that species rank was merited for sundevalli. Although we
can all list many additional data sets required to make a truly informed
decision on this one, the above is all that we have, as far as I can tell.
Based on these data, I'm not really sure what "sundevalli" is,
and I suspect that Payne's idea of multiple colonizations is correct, producing
a population that is an amalgam of various striatus infusions. On the
other hand, if we maintain striatus and virescens as
separate species, then perhaps sundevalli ought to be ranked as a
species also. On the other other hand, the whole
complex needs an overhaul how can striatus and virescens be
kept separate when Old world striatus are continued to be
considered as conspecific?
Nonetheless, I'm intrigued
by the difference in bare-part colors and displays. What I need to know for me
to vote YES on species rank is whether those striatus-like phenotypes
also share those same differences from mainland striatus. At that point I'd lean towards species rank
for sundevalli as a highly variable species.
English name: The
taxon sundevalli is called "Lava Heron" by those authors
who consider it a separate species, including Harris (1982).
Lit Cit:
Harris,
M. P. 1973. The Galápagos avifauna. Condor 75: 265-278.
Harris,
M. P. 1982. A field guide to the birds of the Galapagos. Collins, London.
Kushlan,
J. A. 1983. Pair formation behaviour of the Galapagos
Lava Heron. Wilson Bull. 95: 118-121.
Martínez-Vilata, A. and A. Motis. 1992.
Family Ardeidae (herons). Pp. 376-429 in del Hoyo et al., Handbook of the birds
of the World, Vol. 1. Lynx Edicions, Barcelona.
Payne,
R. B. 1974. Species limits and variation of the New World Green Herons Butorides
virescens and Striated Herons B. striatus. Bull. Brit.
Orn. Club 94: 81-88.
Van
Remsen, 17 May. 2002
________________________________________________________________________________________
Comments from Stotz: “On Lava
Heron, it seems to me that the situation on the Galapagos is likely analogous
to the Slate-colored Coot situation, with a variable population with one form
resembling the near relative (striatus in this case, Fulica americana
in the other). I have to admit that it seems hard for me to imagine that sundevalli
and striatus could be conspecific, if striatus and virescens
are not. It further seems hard to imagine that Old World and New World striatus
are best treated as conspecific. But maybe I am actually arguing here for
the status quo, until somebody figures this all out...
From Nores: SI. Estoy de acuerdo con elevar a Butorides striatus sundevalli
al rango de especie. El color negro y las partes desnudas de la cara celestes
parece que son suficientes."
From Robbins: "YES.
Although the Hayes' article didn't provide any definitive answers, if we are
going to recognize both virescens and striatus as species, then
we should elevate the Lava Heron to species level. Thus, I vote "yes"
for this proposal.
From Stiles:
"Lava Heron a species: A very qualified YES. The argument for raising
(again) virescens to species rank was that the 'intermediate'
buff-necked birds represented variation within striatus, rather
than hybrids. By the same logic, one could say that variation from light to
dark birds on the Galapagos was simply variation within sundevalli,
with the additional point that all have the same distinctive soft part colors.
Given the sedentary nature of striatus, multiple colonizations
doesn´t seem all that likely given the distances involved. I am not aware of
any detailed comparisons of morphology or displays in OW vs. NW striatus. So,
if the AOU recognizes virescens as a species, I think that consistency
demands that we recognize sundevalli as well. If at some point the
AOU reverses itself and 'sinks' virescens, then we reconsider! Un
the meantime, more data on those Galápagos pale birds seems in order.
From Schulenberg:
"What a nightmare.
"I am voting
"NO" on this proposal, largely because I don't feel that I understand
what is going on, and I am reluctant to make any changes if I don't understand
what is going on.
"I have no field
experience with sundevalli. The Field Museum has five specimens: one
juvenile/immature that is "gray", two "black" specimens,
and two "gray" adults. The two gray adults clearly are paler than are
the classic black sundevalli, but also are darker than are most if not
all of our striatus from Ecuador and Peru (and also lack any trace of
the rusty or buff color typically present on the necks of South American striatus).
"Clearly, with only
these few specimens I can't confirm (or refute) Payne's suggestion that plumage
color in Galapagos Butorides spans the full spectrum between gray and
black. It would be nice to know whether indeed that is the case. This is
especially on my mind after I queried two people close at hand who each had
made multiple visits to the Galapagos. Both were of the opinion that all to
almost all of the Butorides that they had seen in the Galapagos
clearly were "black" *or* "gray". (Although their
recollections as to the relative abundances of the two plumages did differ!).
This kind of limited anecdotal data, not supported by specimens, is weak, I
know, but since it seems to fly right into the face of what Payne reported I
feel as if I have to call everything into question: what *is* the prevalence of
the "gray" plumage? Is there intergradation or not? and if so, how
extensive is it? Is there assortative mating? etc.
"I don't think that
recognition of Butorides virescens as a species compels us to
recognize sundevalli. The recognition of virescens as a
species was driven, I thought, not only by the fact that virescens and striatus
are "different", but also that the breeding distributions of virescens and striatus
approach one another in Panama with little evidence of hybridization. It's not
clear to me how that situation would be relevant to whatever is happening on
the Galapagos.
"So, to properly
evaluate the situation on the Galapagos, I think we need to know a lot more
about what several things that we don't know (but that might not be that hard
to get to the bottom of, if someone put their mind to it: Van, can you interest
David Wiedenfeld in pursuing this?).
From Alvaro Jaramillo: "YES,
the plumage and bare part differences (important in pair formation presumably)
between sundevalli and striatus are enough for me to
consider this taxon a separate species. The open
question is what the grey individuals on the Galapagos are, grey morph of sundevalli
or real striatus that have recently colonized the islands.”