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BIOLOGIA
CENTRALI-AMERICANA.

ZOOLOGY, BOTANY,
AND

ARCH AOLOGY.

EDITED BY

FREDERICK DUCANE GODMAN, DOL, FRS,,

AND

OSBERT SALVIN, MA, FERS.

INTRODUCTORY VOLUME.

BY

FREDERICK DUCANE GODMAN, D.C.L., F.R.S.

1915,

To My Beloved Wife

ALICE MARY GODMAN
who has taken the deepest interest

and given me
much assistance and sympathy
an the completion of
this work

a2

*

CONTENTS.

PREFACE

Puatss J., IT.

INTRODUCTION. . . .
Physical Features, etc., of the Area treated .

Sources from whence our Material was obtained

Itinerary of Mr. G. C. Champion’s Travels in Central America, 1879-1883

List of Completed Volumes, Zoology, Botany, and Archeology
Analysis of Contents of each Volume
Summary of Contents of each Volume .

Origin, etc., of the Fauna and Flora of Central America :—
Fauna: .
Mamata. By R. I. Pocock rr ree
Reptitia, Batracuis, and Pisces. By C. Tate Regan .
AracunipA (Opiliones and Acari excepted). By R. I. Pocock

Cuitopopa, Dieroropa, and Protorracnzata. By R. I. Pocock .

Frora. By W. B. Hemsley
Mars I.-VIII.

87, 142
. 105
118, 142
. 183
~ 145

CORRIGENDUM.

Page 55, line 11 from top. For Protobracheata read Prototracheata.

PREFACE.

I FreeL that an apology, as well as an explanation, is due to the subscribers of the
‘Biologia Centrali-Americana’ for the length of time that has been occupied in
the production of this work. When it was commenced, in September 1879, it was
estimated in our prospectus that, when completed, the Zoology would ‘not much
exceed sixty parts, equivalent to about twelve volumes of 500 pages each,” and that
twenty parts would suffice for the Botany, the two subjects to be issued concurrently.
The inclusion of the Archeology was not at that time contemplated, and this subject
was only undertaken later in consequence of the investigations made by my friend
Mr, A. P. Maudslay of the famous ruins in Central America, which, together with his
beautiful photographs, made a valuable addition to our knowledge of the country.
We had, moreover, underestimated the vast amount of additional material which
subsequently came into our possession, and thus necessitated the extension of the

work to a total of 215 parts, or 63 volumes,

It had been our intention at the termination of the work, and after a careful study
of the Zoological and Botanical material accumulated from this hitherto little-known
but exceedingly rich ‘country, to have summarised the result and discussed its bearing
on the interesting subject of geographical distribution, Salvin’s death after a long
illness, and my own advancing years and ill-health, compelled me to abandon this
project, and I should have been obliged to content myself with the conclusions arrived
at by the various contributors in their respective Introductions had it not been for the
assistance of Messrs. R. I. Pocock and C. Tate Regan, to whom I am greatly indebted for
their respective articles on the Mammals, Reptiles, Fishes, Arachnida, Chilopoda, etc.,
which are included in the Indroductory Volume. As regards the Insecta generally,
which occupy such a large portion of the work, so little is as yet known of the fauna

of other tropical regions that no satisfactory comparison can be made.

vill PREFACE.

Mr. W. B. Hemsley, who had previously contributed the volumes on the Botany, had
almost completed an article on the geographical distribution of the Flora, practically
bringing this subject up to date, when, I regret to state, his health completely failed,
and he was reluctantly obliged to relinquish his task. Recently he has, however,
been able to furnish me with a précis of his conclusions, which forms a valuable

addition to our knowledge of the subject.

It now only remains for me to offer my grateful acknowledgments to all those
who have assisted me with their various contributions, and without whom the work
could not possibly have been undertaken. To my Secretary, Mr. G. C. Champion,
I am specially indebted for the valuable assistance he has rendered as collector,
contributor, and also as subeditor, in which last capacity his advice has been of
inestimable value. His knowledge of Entomology, especially of Coleoptera and
Rhynchota, has made him one of our most important contributors, and he has
either undertaken alone, or shared in the production of, no less than nine volumes
of the ‘ Biologia.” My warm thanks are also due to my assistant, Mr. A. Cant, who
has given very important help during the progress of the work, both in setting the
insects, in labelling and arranging them, as well as in making very careful dissections

aud preparing slides for microscopic examination.

F. D. G.
June 1915,

PLATE I.

LQ. Godesti-

PLATE IL

INTRODUCTION.

A sHorT account of the events that led to the publication of the ‘ Biologia Centrali-
_ Americana’ may be of interest to our readers, and I will therefore first give a sketch
of the early days of Salvin and myself, so far as they have a definite bearing on the
study-of Natural History, and of the circumstances which drew our attention especially
to Tropical America.

Osbert, the second son of Mr. Anthony Salvin, the eminent architect, was born at
Finchley in 1835 and educated at Westminster and Cambridge. That he developed
a very early taste for natural history is clear from the series of bird skins, now in
the Natural History Museum, collected by him as a boy and labelled ‘ Finchley.’

I, Frederick DuCane Godman, third son of Joseph Godman, of Park Hatch, Surrey,
was born in January 1834, and at the age of ten went to Eton, but three years later
a very severe attack of what was then called low fever necessitated my removal,
and for some years I was unable to work at all. When my health was sufficiently
re-established, I received instruction from tutors until I was eighteen years old, when
I made a trip to the Mediterranean and Black Sea, visiting Gibraltar, Southern Spain,
Athens, and Constantinople en route. During the time spent at home I interested
myself in Natural History, paying special attention to the British Mosses and Ferns,
of which I made a considerable ccllection. Birds were always a source of delight
to me, and I could recognise a large number of British species as well by their flight
as by their note. |

In 1853 I entered Trinity College, Cambridge, as an undergraduate, and Salvin, in
the following year, went to ‘Trinity Hall, of which College he became a scholar; he
graduated as a Senior Optime in the Mathematical Tripos and was afterwards made
an Honorary Fellow. With similar tastes, it was only natural that we soon met
and became fast friends, thus forming that close intimacy which only terminated
with Salvin’s death on June Ist, 1898. Salvin was a skilful mechanic, and very
ingenious in carpentry and cabinet making. Whilst still at Westminster, with the
assistance of his elder brother, he built a boat thirty feet long and fitted it with
a steam engine, the whole of which, with the exception of the boiler, was made
by the two brothers. This boat was launched on the river, and in it they went to
a Thames regatta, but, having tested its powers and proved its capability, their object

BIOL, VENTR.-AMER., Introd. Vol., January 1915. B

2 INTRODUCTION.

was achieved, and the boat was finally sold. Some years afterwards, at Duefias, in
Guatemala, when we required specimens of the duck and waterfowl which frequented
the neighbouring lake, Salvin again turned his hand to boat building. ‘This time the
ribs and frame were made of sticks of green wood cut and fastened together ; over
this, the hair having previously been removed, a raw ox-hide was drawn. and as the
hide shrank, it bound the whole tightly together and made an excellent boat, easily
accommodating two people. In this craft we had many sails upon the lake and
obtained examples of the birds resorting there.

During our College days, Salvin and I made frequent expeditions together to the
fens and other places in the neighbourhood of Cambridge, in order to collect birds’
eggs and lepidoptera. On one occasion we heard of a bustard which had been seen
in Wicken Fen, and we spent a couple of days searching for it, but with no intention
of shooting so rare a visitor. We found both its foot-tracks and some shed feathers,
but, as we learned afterwards, the bird had been shot at and probably wounded by
one of the fenmen, as it was never seen again. We also spent our leisure hours in
Baker’s shop, the well known bird stuffer in the Trumpington Road, skinning and
setting up birds—an experience which we found of great service to us afterwards when
in Central America and on other expeditions,

A good deal of fenland being then undrained, Swallow-tailed butterflies were always
to be found, and we collected the larvee and bred them in Baker's shop. ‘The
‘Large Copper’ had so recently become extinct, that we searched in vain for it, though
Brown, the tailor in Cambridge, who was an ardent British lepidopterist, had a long
series in his cabinet, mostly specimens bred from the larve he had collected a few
years previously.

While still at Cambridge there were several other University men keenly interested
in Ornithology, notably the two brothers Newton, Simpson, and my brother Percy, and
after our spring rambles we used to meet in each other's rooms and discuss the result
of our various expeditions. It was at one of these meetings in 1807 that it was first
suggested that some record should be kept of these proceedings, and the idea of
estabiishing a Magazine solely devoted to Ornithology was mooted, but nothing
further was done till November 17th in the following year, when a meeting took
place in Alfred Newton’s rooms in Magdalene College, at which Salvin and myself,
Simpson, Wolley, Sclater, Newton, and other ornithologists were present. Before
the party broke up it was resolved: ‘That an Ornithological Union of twenty
members should be formed, with the object of establishing a new Journal devoted to
Birds: that Lieut.-Colonel H. M. Drummond should be President, Professor Newton
the Secretary of the Union, and P. L. Sclater should edit the Journal: that the title
of the Journal should be ‘'The Ibis.’ ”

The first volume of ‘'The Ibis’ appeared in 1859, and the Magazine has now reached
its 56th volume, and the Union has over four hundred and forty members.

INTRODUCTION. 3

In 1857, Salvin made a birds’-nesting expedition with the Rev. H. B. Tristram and
Mr. W. EH. Simpson (afterwards Huddleston) through Tunisia and Eastern Algeria, in
which I was to have joined them, but an accident in the hunting-field laid me up
for some weeks and prevented me from accompanying them. ‘The result of this five
months’ journey forms the subject of two valuable papers, one by Salvin, the other by
Tristram, published in the first volume of ‘The Ibis.’ Later in the year, when I had
sufficiently recovered from my accident, I went with my brother Percy to Bodé, in
the north of Norway ; there we remained for some weeks exploring the surrounding
country and were fortunate enough to meet with and secure the eggs of the Great
Snipe (Scolopax gallinago). Taking the steamer northward to the Alten River, we
crossed Lapland on foot to Haparanda, on the Gulf of Bothnia, paying John Wolley a
short visit at Muonioniska. Before returning home we visited Stockholm, St. Peters-
burg, and Nijnei-Novgorod. A short paper on the birds obtained on this journey
appeared in ‘The Ibis’ for 1861.

In the autumn of 1857 Salvin paid his first visit to Central America, in company
with Mr. George Ure Skinner, a gentleman well known to both Botanists and
Ornithologists through the collections of crchids and birds he had brought from that
country on previous expeditions. Salvin undertook the journey, at the request of
Messrs. Price & Co., to examine and report upon the nuts of a palm which it was
thought might be used in the manufacture of candles. ‘The palm-nuts, however,
proved to be useless for practical purposes, and Salvin spent the remainder of his time
in travelling through the country and making a collection of birds and insects. He
reached Belize, British Honduras, in December 1857, and after spending a few days
there, proceeded down the coast to Yzabal and thence by easy stages to Guatemala City,
making Duenas, 3U miles south-west of the capital, his headquarters for six months.
Salvin made two excursions to the Pacific coast region and one to the Lake of Atitlan
in the ‘Altos.’ Leaving the country towards the end of June 1898, he returned to
England vid San José and Panama. On his return he published a paper in ‘ The Ibis,’
in conjunction with Mr. P. L. Sclater, on the Ornithology of Central America (not
including Mexico), in which the authors enumerated 381 birds, all that were then
known to inhabit that country.

What he saw, however, on this expedition so whetted his appetite that he returned
again to Guatemala in the spring of 1859, with the sole object of studying Natural
History. He revisited Duefias, and collected in the neighbourhood for some months.
In October he went to San Gerdénimo, Coban, and other places in Vera Paz, returning
to Duefias about the end of the year. In March 1860, he was again in Aita Vera
Paz, at Coban, Lanquin, &c., and left for home, wd Belize, in the following month.
On this occasion he added very considerably to his collection of birds, as well as
insects, and as a result wrote various papers in ‘The Ibis’ for 1860 on his
discoveries.

B2

4 INTRODUCTION.

In August 1861, I joined Salvin on his third expedition to Guatemala, and, after
spending three weeks in Jamaica en route, we landed at Belize; thence taking our
passage in a coasting schooner we arrived at Yzabal on the Golfo Dolce. Here we
remained a few days, making preparations for our journey and engaging Indians and
mules to transport ourselves and our baggage to the interior. This place will,
however, always be associated in my mind with my first sight of a living example
of one of the most striking and gorgeous of all butterflies, Morpho peleides. Iwas
sitting on the trunk of a fallen tree in the forest, when it came floating past me, but
1 was so overcome with astonishment and delight at this wonderful vision that,
although I had a butterfly net in my hand, I was utterly unable to rise in pursuit
until it was too late to capture it.

Crossing the Mico range of mountains, we spent a few days at Quirigua, where ©
I first encountered the great Howling Monkey (J/ycetes villosus), which frequents
the dense forest in troops, making night hideous with its howls, which could
be heard in the stillness for a distance of some miles. As we were sleeping in
hammocks swung from the boughs of trees, we were somewhat disturbed in our
slumbers. We also spent our time photographing the old Indian ruins and exploring
the forest in the vicinity. Dry and highly sensitive plates, such as are in ordinary
use now, did not then exist, and every photographer was obliged to carry about the
necessary materials for preparing and developing his own plates, which might either
be wet, entailing immediate development, or dry (tanning), when they could be kept
for some days. The whole photographic apparatus, including chemicals, fitted into
a case, which formed a load for one of our Indian carriers. Our first essay at photo-
eraphy in the forest was not a success, as we found that after exposing the plates
for twenty minutes no details were to be seen. ‘This we discovered was owing to
the dense green foliage overhead, through which the light had scarcely any effect in
dissolving the nitrate of silver on the plate, and consequently no image was produced.
In order to overcome this difficulty, we then hired Indians to cut down the trees which
shaded the objects we wished to photograph. This delayed us a few days, which,
however, we employed in collecting birds and insects, until a sufficient number of
trees were felled to admit light upon the ruins, when we again proceeded to take
photographs of the large monoliths, now obtaining very successful results. These
ruins are fully described and illustrated in the ‘Archeology’ of the ‘ Biologia,’ by
A. P. Maudslay.

From Quirigua we again took the mule track, for it could hardly be called a road,
through the valley of the Motagua River to Zacapa, and thence to Guatemala City.
We spent a day or two at the Capital and then proceeded to Dueftas, where we
remained for some weeks in most delightful quarters at the house of Mr. William
Wyld, a friend of Salvin’s. Our time at this place was devoted to collecting, chiefly
in the high forests of the Volcan de Fuego, the peak of which we ascended, and

INTRODUCTION. . 5

I made a separate expedition to Escuintla in the Pacific Coast region. After our stay
at Duejias we retraced our steps to the Capital, and, crossing the Chuacus Range into
the plain of Salama, we took up our abode at the Hacienda of San Geronimo. Here
we resided for some weeks, finding several species of birds and insects which we had not
previously obtained. From San Gerdnimo we went to Coban, and, after spending some
time collecting in the neighbourhood, we visited Cubilguitz and Choctum in the low
damp forest of Alta Vera Paz. At Cubilguitz, unfortunately, I contracted a sharp
attack of fever, which obliged me to remain for some days at Coban to recruit and
prevented my accompanying Salvin on his long and arduous Journey on foot to Peten.

When I had recovered sufficiently I returned to San Gerénimo and then went to
Buenaventura on the upper waters of the Motagua River, there called the Rio Grande,
where I employed Indians to poison some nine miles of the water in order to make a
collection of the fish. Before commencing operations I noticed one of the ‘ mozos’
lying flat beside the river, wafting some burning material over the surface of the water,
and, upon questioning him, I elicited that he was propitiating the spirit of the river
in order that success might attend his efforts and the fish be permitted to die. The
method adopted for this purpose was to beat the plant (Tephrosia toxicaria, Pers.*,
B. C.-Am., Bot. i. p. 258) on the rocks until a froth not unlike soap-suds was formed,
this when mixed with the water caused the fish ‘to sicken and come to the
surface. At intervals V-shaped wicker guides were placed, so that the fish floated
down to the point of the V, where they were then collected in baskets, and when not
otherwise required were used as food by the Indians, who considered them perfectly
wholesome. Vast numbers were thus obtained, and from them I made a selection and
preserved a good many specimens in spirit (aguardiente), but was somewhat disappointed
to find there were but few species represented. On my return to the Capital I journeyed
to the Alotepeque silver-mines in company with the manager, and thence to Copan,
Honduras, where, after spending a couple of days in examining the interesting ruins, I
proceeded vid Zacapa to Yzabal, and there met Salvin on his way back from Belize.
Here we again parted, I returning to England, while he started for Duefias and the
interior, passing through the ‘Altos,’ staying at Totonicapam and Quezaltenango,
and making expeditions to the Costa Grande, Retalhuleu, and the lagoons of the
Pacific coast at Huamuchal, close to the Mexican frontier. Salvin returned home
early in 1863. .

During our outward journey to Guatemala in August 1861 Salvin and I passed
through the Azorean Archipelago, and I then wished I could stop and explore these
islands, but onward we went and soon again lost sight of land. I frequently
throughout the voyage recalled these isolated islands and determined, should
opportunity occur, that I would explore their fauna and flora at some future date.

* Order Leguminose.

6 INTRODUCTION.

It was not, however, till the spring of 1865 that I was able to carry out this project.
The careful researches of Wollaston and others had brought to light many interesting
forms from Madeira, the Canaries, and Cape Verde Islands; but the Azores had been
very imperfectly explored, and it was with the idea of giving a more satisfactory
account of the natural history of these islands, and to trace their relationship to the
neighbouring Archipelagos, that I decided to investigate their fauna. Oranges from
St. Michael’s then formed almost the only trade with England, and in connection with
the business large numbers of schooners were employed, but with this exception
there was no regular, direct communication with England. <A small steamer, which
carried the mails, left Lisbon once a month for the two nearest groups of islands, but
rarely visited the two outer ones. Accompanied by my brother, Captain Temple
Godman, I took a passage in this vessel, and shortly after our arrival at St. Michael’s
we were joined by Mr. Brewer, a well known coleopterist whom I had engaged for the
purpose of collecting. Interest in island faunas had been much stimulated by the
publication of Darwin’s “ Geology of the Voyage of the ‘ Beagle’” and other works on
the subject. The question was whether the Azores had in former times formed part
of a continent now submerged, as Professor Edward Forbes believed, or whether they
had been thrown up from the sea bottom by volcanic agency. After spending four
months on the islands, during which time I visited all except Santa Maria, I came to
the conclusion that they had always been volcanic islands, and that they derived their
fauna and flora from neighbouring lands. In 1870 I published a small octavo book
entitled ‘The Azores,’ enumerating the plants and animals as far as then known, and
setting forth my reasons for the conclusion above stated.

The visit to the Azores was followed in 1873 by an expedition to Madeira and the
Canaries, in order to compare their respective faunas more critically, but unfortunately
the regulations in respect to quarantine were then so stringent that, apart from Madeira,
my investigations were limited to the island of Teneriffe. There can, I think, be no
doubt that the conclusion I had already formed with regard to the Azores was fully
borne out in these islands also.

In the meantime, Salvin was residing at Rotherham, Yorkshire, looking after some
ironworks in which he was financially interested; but this was not a congenial employ-
ment, and he soon gave it up.

In 1865 he married Caroline, daughter of J. Whitaker Maitland, of Loughton Hall
in Essex, and they lived for some years at 23 The Boltons, South Kensington, which
became for a time the headquarters of our Museum.

Although both Salvin and I had jointly collected ever since our undergraduate days, it
was not until the material was housed at 8. Kensington that we really did serious work
together ; but from thence onwards we spent the greater part of the week in London
arranging our collections, publishing papers on them, and attending the meetings of
various scientific societies of which we were both members.

INTRODUCTION, 7

Salvin’s fourth and last visit * to Guatemala was made in conipany with his wife, to
whose skilful brush we are indebted for the coloured plates of the plants figured in the
Botany of the ‘ Biologia.’

They sailed in a Royal Mail Steamer in April 1873, touching at St. Thomas and
Jamaica; then crossing the Isthmus of Panama they reached the City of Guatemala early
in June. Proceeding thence to Duefias, which became their headquarters for some
months, Salvin occupied himself in collecting in the forests on the mountain slopes.
Together they ascended the crater of the Volcan de Fuego, and a few days later that
of Acatenango. Leaving Duefias for Atitlan they made the ascent of the peak from
Santa Lucia on Jan. 17th. Subsequently they visited Mazatenango, the coffee estate
of Las Nubes (Cerro Zunil), Quezaltenango, the Lake of Atitlan, Solola, Pantaleon, and
San Gerdnimo, and after a short stay at the last named place the journey was continued
to Coban. Having revisited the Capital they left Guatemala in March 1874. Again
erossing the Isthmus of Panama, they sailed for the United States and visited the
museums of New York, Boston, Philadelphia, and Washington, and made the acquain-
tance of the leading scientific people, returning to England on June 4th, 1875.

In the autumn of the same year, Salvin was appointed to the Curatorship of the
Strickland Collection of Birds at Cambridge, which necessitated his residence at the
University. On his giving up the house in South Kensington in 1873, it had become
necessary to find fresh quarters for our museum and library, which now occupied a
considerable amount of space, and we took for this purpose a house in Tenterden
Street, Hanover Square, to which they were transferred. ‘The building being rather
larger than we required, we shared it with some of our ornithological friends, of whom
Lord Lilford, Dresser, and Seebohm were among the number. After the evening
scientific meetings of the Zoological Society, the offices of which were then in Hanover
Square, our rooms became a favourite social resort of ornithologists, and many
pleasant and instructive hours were spent there.

During his stay at Cambridge, Salvin came frequently to Tenterden Street and
worked at the collections with me, and continued so doing until the death of his father
in 1880, when he succeeded to his property at Fernhurst, Sussex, where he afterwards
resided, continuing, however, his work in London as before. In the autumn of 1878
we moved our museum and library to 10 Chandos Street, Cavendish Square, and
here they remained till after Salvin’s death. In 1907 the house at Chandos Street
was given up and the library transferred to 45 Pont Street, S.W., while the collections
still remaining in our possession were handed over to the British Museum.

In addition to the material obtained during our various visits to Central America
and that sent us by the natives we had trained, we found it necessary, for the sake of
comparison, to acquire a more thorough knowledge of the South American fauna, and,

* In the notice of his journeys given in the Introduction to the ‘ Aves’ (p. vili) the second expedition,
1859-1860, was not mentioned, and the last visit was incorrectly stated to have been made in 1867.

8 INTRODUCTION.

with that view, employed various expert collectors, whose names are recorded in the
body of the work, to visit special localities in Colombia, Ecuador, Peru, Guiana, and
other places in South America. We continued to receive consignments from them for
several years without having any idea of publishing a connected account of the results,
and a very large amount of material, especially amongst the birds and insects, was thus
accumulated. It was not till the year 1876 that it was suggested that the ‘ Biologia ’
should be undertaken, and three years later (September 1879) the first part appeared.

In the meantime, various collectors were sent by us to Guatemala and other parts of
Central America.

Our method of publication was to bring out six quarto parts a year ; each part to
contain twelve sheets made up of various subjects with six coloured plates, the plates and
letterpress so numbered and paged that the parts might ultimately be broken up and
bound together in their respective volumes when completed. In this way it was possible
to keep several subjects in progress at once, and the plan answered well. We were,
however, unable to adhere to the original scheme of completing the work in 60 parts,
owing to the ever increasing amount of material received from our collectors—an
amount so great that 215 Parts of Zoology alone have been required, the dates of
issue extending over a period of 36 years. Even now some families of Insects, the
Crustacea, &c., have not been dealt with—though this is chiefly attributable to the
fact that no experts on these subjects were available.

On arrival in England, the various consignments were opened, every specimen
labelled with its exact locality, and the name of the collector attached. The animals
sent were then sorted into their respective orders and families, and as occasion offered,
handed over to specialists to be worked out. It was obviously impossible that we
could undertake every subject, but the birds and the butterflies we set aside for our
own share of the work. The names of the various authors who kindly helped us will
be a sufficient indication of our good fortune in securing the services of so many
eminent men, all of whom joined the enterprise with great spirit.

In 1886, in company with Mr. H. J. Elwes, I visited Bombay, Simla, Delhi, Benares,
Calcutta, and Darjeeling; thence crossing the Rungeet River by a swinging bamboo
suspension bridge we entered native Sikkim, collecting plants, birds, and butterflies
en route. At Darjeeling I purchased a large collection of butterflies, made by
Lidderdale, which at the time was considered very representative of the Northern Indian
fauna. Starting again from Darjeeling we made a second trip, this time on horse-
back, travelling by the Government road constructed by Sir Richard Temple along
the Nepaul ridge till we reached Falute (16,000 ft.), and there passed the night in
a fairly comfortable bungalow. Next morning we witnessed a magnificent sunrise and
obtained splendid views of Mounts Everest and Kinchinjunga rising from the mighty
Himalayan range. I also visited Madras and Southern India and Ceylon before
returning home.

INTRODUCTION. 9

In the autumn of 1887, having been ordered abroad for the benefit of my health,
I decided to visit Mexico, as in working out the Central American fauna, especially
the Birds, we found ourselves more deficient in material from that country than from
further south. Crossing the Atlantic to New York I took the train to Mexico City,
then a six days’ journey, entering the Mexican Republic at El Paso (3700 ft.) on the
Rio Grande. From this river the land gradually rises and spreads out into the great
plateau of Central Northern Mexico, bounded on the east and west respectively by
ridges of high mountains covered with pines, and falling abruptly on the east to the
Atlantic and on the west to the Pacific. The rainfall being chiefly on the two coasts,
the plateau is extremely dry and arid, and the vegetation consists largely of cacti,
yuccas, agaves, and mesquite (Prosopis), with willows and poplars along the margins.
of the few water-courses or lagoons. The early part of the journey was passed during
the night, but next morning it was evident that a considerably higher elevation had
been attained, and on reaching Zacatecas (8000 ft.), a large mining district, the aspect
of the country was very desolate and unpromising for natural history purposes, and
continued much the same for some distance, but improved a little nearer to the Capital.
After spending a few days in Mexico City in making necessary preparations, I was
joined by W. B. Richardson, an American bird-collector, and taking him with me
started for the Atlantic coast. The first part of the journey, still on the high plateau,
was through fields of ‘agave’ grown for the production of the fermented drink
called ‘ pulque,’ so much beloved by the Indians. In about six hours we reached
Esperanza, in the State of Puebla, at the Eastern edge of the plateau ; here the train
enters the wonderful gorge by which it descends to Orizaba, Cordova, Atoyac, and
Vera Cruz. The scenery at once changes, the vegetation becoming luxuriant on
entering the region of the rainfall.

Before reaching Orizaba we crossed and recrossed the gorge by a series of viaducts
amongst palms, tree ferns, and tropical plants, loaded with orchids and tillandsias. At
Orizaba we spent a few days coilecting, and I was there joined by Mr. and Mrs. H. H.
Smith, who went with me as far as Atoyac, where they remained for some time before
crossing to the Western side of Mexico. The Smiths had previously been in Brazil,
where they made large collections of insects, which are now in the Pittsburg Museum.
Mrs. Smith was also skilled in skinning birds which were shot and brought to us by
the Indians, and through her we made many additions to our store of ornithological
treasures. Leaving the Smiths at Atoyac, a village at the foot of the steep descent from
the plateau—still, however, about 1500 feet above the sea and about fifty miles from
the coast,—Richardson and I continued our journey to Vera Cruz, the land gradually |
sloping down to the Atlantic and forming a savanna or plain of sandy ground, sparsely
covered with grass and scrub. After spending a few days in collecting, chiefly to
the north of the town, we took the mule tram to Jalapa, which being on higher ground
reaps the benefit of the rainfall and the vegetation is far more luxuriant. At this

BIOL. CENTR.-AMER., Introd. Vol., January 1915. C

10 INTRODUCTION.

place I engaged Mateo Trujillo, a half-breed Indian, who accompanied me during the
greater part of the time I was in Mexico and proved a very skilful collector. He was
a first rate climber, and amongst other things made a considerable collection of the
frogs, newts, and insects which inhabit epiphytical Bromelias growing on the trees in
the neighbourhood of Jalapa. We made an excursion to Misantla and Papantla, on
the low ground near the coast at the foot of the mountain range, which proved to be
very good collecting ground and added considerably to our birds and insects. On
leaving Misantla we returned to Jalapa and over the Cofre de Perote to Esperanza, and
thence by train to Mexico City. From the Capital I made two expeditions, the first
in company with Mr. Flohr to the pretty town of Cuernavaca, and thence to the caves
of Cacahuimilpa, where he hoped to have found some blind insects. The second and
also interesting expedition was to Morelia and Lake Patzcuaro, where I added to the
collection of Birds, but it was attended with no very valuable results.

On returning to the Capital I next visited Yucatan, crossing the Gulf of Campeche
from Vera Cruz to Progreso by steamer, thence to Merida by train, where the railway
then ended. My first object was to visit the well known naturalist Dr. Gaumer, who
kindly accompanied me to the celebrated ruins of Ticul and Uxmal, a distance from
Merida of about forty miles, which journey was performed on horseback. The low
forest through which we passed is said to be still frequented by the beautiful ‘ Pavo
real’ (Meleagris oceliata), but it has become so rare in the neighbourhood that we
ourselves did not meet with it. Leaving Yucatan I went back to Mexico City, where
I was joined by Mr. and Mrs. Elwes, and together we went to Jalapa, thence riding
across the country to Cordova, a most delightful journey of three days, and obtaining
magnificent views of the Volcano of Orizaba, with its snow-clad peak. We continued
our journey to Puebla and Mexico City, and thence to Amecameca at the foot of the
volcanoes Iztaccihuatl and Popocatepetl; the latter we ascended as far as the pine belt,
shooting many interesting birds and collecting plants and insects. We left Mexico in
the spring of 1858, having been absent from home about five months, and returned to
England wd California and New York.

For several years after my return from Mexico, Salvin and I continued diligently
to work out the material on hand and the ever increasing amount sent over by our
collectors. Salvin’s failing health finally obliged him to relax his efforts, and though
he still came to London as formerly, he was unable to take the same active part
in the work and the difficulty of concentrating his attention on any one subject
became increasingly great. He died suddenly at Hawksfold, Fernhurst, Sussex,
Jure Ist, 1895, 'eiving me alone to complete the ‘ Biologia.’

The severance of a friendship such as ours had been for forty-four years was a terrible
blow to me, for we were more intimately connected than most brothers, and, besides
the personal loss, I missed his knowledge and experience in all things connected with
our book. At the time of Salvin’s death, 14] Parts of Zoology (completing 13 volumes),

INTRODUCTION. 11

the whole of the Botany (5 volumes), and nine Parts of Archeology had been issued :
since then 74 Parts (142-215) of Zoology, completing 39 more volumes, and 8 Parts
of Archeology have been required, to bring these subjects to aconclusion. It
was with a heavy heart that I took up my pen again. With the assistance of
Mr. G. C. Champion I continued Vol. II. of the Rhopalocera and finished it in 1901,
and in 1904 Vol. III. of the ‘ Aves’ with the help of Dr. R. B. Sharpe, of the British
Museum, was also completed.

Salvin had made a special study of the Procellariide, a very difficult group of birds,
and for a long time we had missed no opportunity of adding specimens of Petrels to
our collection. After writing the catalogue of this family for the British Museum, he
intended to publish an illustrated monograph on the subject, and for that purpose
40 plates had already been drawn by Keulemans. It remained for me to carry out
his intention, and having again sought the aid of Dr. R. B. Sharpe we completed
the work in 1910, enumerating 123 species, and illustrating them with 106 coloured
plates.

It will be seen that the travels of Salvin and myself covered only a comparatively
small part of Mexico and Central America, but we employed a considerable number
of expert collectors to travel in districts we had not visited, and they continued to send
us the results of their labours for some years after we had left. Amongst them
must specially be mentioned W. H. Richardson, Mr. H. H. Smith, and Mateo Trujillo,
all of whom accompanied me while I was in Mexico, and Lloyd and Armstrong, who
devoted their attention particularly to the Northern provinces of that country.

In working out the Mexican Birds we found ourselves hampered for want of
an authentically named collection of North American species for comparison, which
did not exist in Europe at that time. In order to remedy this, I acquired the Henshaw
collection, numbering 13,326 specimens, and this was rendered still more valuable
through the courtesy of the authorities at the United States National Museum,
who allowed Mr. Ridgway, the highest authority in America, to go through and
verify all the names on the labels attached. I likewise purchased 2500 carefully
named birds from Florida from Mr. W. E. D. Scott, 321 named specimens from
California, Texas, &c., from Mr. C. K. Worthen, and a series of Mexican birds from
Sefior F. Ferrari-Perez ; and these collections proved of great advantage to us.

In a similar manner we dealt with the insects, &c. I bought H. W. Bates’s
collection of butterflies, including those he obtained from the Amazons, as well as
that of Herbert Druce, containing the ‘ Kaden’ types; the first set of the extensive
series of Mexican and Central American Coleoptera amassed by A. Sallé (including the
types of several of the older authors, and some thousands of specimens found by
himself or by M. Boucard). I also acquired the general collection of Heteromera of
F. Bates (22,390 specimens); a portion of Dr. J. S. Baly’s collection of Phytophaga ;
the Janson collection of Elateride (25,000 specimens); various Reptilia, Coleoptera, &c.,

C2

12 _ INTRODUCTION.

collected by A. Forrer in N.W. Mexico and the Tres Marias Islands; a second set of
the very large number of Coleoptera obtained by C. T. Hége in his two expeditions
to Mexico, the ‘first set now in the Berlin Museum, having been retained by his
employer, Mr. Flohr ; collections of insects from Messrs. Becker, Biolley, Blancaneaux,
Conradt, Gaumer, Janson, Lankester, Morrison, Staudinger, Underwood, Van Patten,
Wittkugel, &c. In addition to this material, we had, of course, the whole of that
procured by our other collectors, E. Arcé, G. C. Champion, and H. Rogers. Further
details are appended on pp. 44, 45.

All the insects from Mexico and Central America, the Sallé and Janson collections
of beetles, our own general collections of birds and butterflies, and the Henshaw
collection of birds, have been presented by us to the British Museum, and are being
gradually incorporated with the National Collection.

The various accessions are enumerated in detail in Vol. II. of the ‘ History of the
Collections contained in the Natural History Departments of the British Museum’
(1906) and in the subsequent Annual Reports of that Institution. The first instalment
of Neotropical birds (50,120 specimens) was presented in January 1885, and other
instalments followed from time to time till the whole of them became the property
of the Nation. Amongst the insects, up to 1906, the total number of specimens
given in the ‘History’ is as follows: Coleoptera (85,920), Lepidoptera Rhopalocera
(17,829), Lepidoptera Heterocera (12,883), Diptera (17,525), Hymenoptera (10,004),
Rhynchota Heteroptera (5543), &c. These figures do not include the Rhynchophora
or weevils (22,793), the Staphylinide and water-beetles (9474), the Odonata (3000),
‘the Rhynchota Homoptera (5509), the supplementary unworked parasitic Hymenoptera
(6293), &c. From 1906 onwards the remaining collections have been handed over
to the Museum as soon as the enumeration of the species was completed; that of
the Coleoptera was finished in 1911. Our own general collection of butterflies
probably included nearly 100,000 specimens, and the beetles alone from Mexico and
Central America perhaps double that number. Besides these a considerable number
of mammals, reptiles, fish, &c., of which no account was kept, were presented to the
National Museum.

F. D. G.

beddg

PHYSICAL FEATURES ETC. OF THE AREA TREATED.

Tue area treated in the ‘Biologia’ includes the whole of Mexico south of the
Rio Grande as far as El Paso, thence to the Gila River, and following it as far as
the Gulf of California (but excluding Baja or Lower California). The distant
Revillagigedo Islands have been added, owing to certain species of sea birds being
common to these islands and the Tres Marias on the western coast of Mexico. Further
south we include British Honduras, Guatemala, Salvador, Honduras, Nicaragua, Costa
Rica, and Panama, which collectively are spoken of as Central America. The country
stretches in a south-easterly direction, having a width in the north of about 1140 statute
miles and gradually decreasing till at the Isthmus of Panama it does not exceed
45 miles, and the land only attains an altitude of 300 feet (tierra caliente). In
shape Mexico and Ceritral America have been likened to a cornucopia, which
collectively they somewhat resemble. Of the physical aspect of each country I
propose giving an outline, but as the travels of Salvin and myself covered only a small
portion of the area, I shall supplement the account with extracts from other writers ;
moreover, upwards of fifty years have elapsed since I was in Guatemala, and doubtless
great changes have taken place in the interval throughout the whole of Mexico and
Central America, partly owing to the extensive destruction of forest for the purpose
of cultivation, the construction of railways, and the wanton devastation of large tracts
by fire. ‘Thus various places which in our time were good collecting grounds are no
longer so, and many species of both animals and plants must either have migrated
or become extinct. Earthquakes, too, have done much to alter the configuration of
the land, as well as the nature of the vegetation.

The country is divided by the natives into three zones—the ‘ tierra caliente,’ ‘ tierra
templada,’ and ‘tierra fria’ (or hot, temperate, and cold climates respectively). The
tierra templada corresponds on an average with an elevation of 3000 to 5000 feet, but
the natives of the Mexican State of Vera Cruz draw this imaginary line at a level very
different from that used by the people on the western slope of Mexico, For instance,
Chilpancingo at 4000 feet in Guerrero has no tropical vegetation and the climate is
much cooler than in the State of Orizaba at an almost similar altitude on the eastern
slope, which is in every sense subtropical.

The difference is due to the greater rainfall on the Atlantic coast, for the prevailing
wind in passing over the ocean is charged with humid air, and when driven by the
mountain ranges into a higher and colder elevation, the moisture is then precipitated
as rain, thus inducing a much more vigorous vegetation than is found on the Pacific
slope. |

14 PHYSICAL FEATURES ETC.

MEXICO.

The general aspect of the Mexican plateau, as far south as the City of Mexico, has
been previously alluded to in the account of my journey there in 1887-1888 (antea,
pp. 9, 10). On each coast, between the mountain ranges and the sea, there is a com-
paratively narrow strip of low land, producing a tropical vegetation, which on the
Atlantic extends a little to the north of Tam pico, where the forests with epiphytical
orchids suddenly disappear; while on the Pacific the low land terminates a little above
Mazatlan, where the coco-palm ceases to grow, but the vegetation generally is less
luxuriant on this coast than on the Atlantic. The plateau itself is arid in consequence
of the rainfall being precipitated on the two coastal ranges, and produces but a scanty
vegetation, consisting chiefly of Yuccas, Agaves and Cactacee, and Mesquite (Prosopis).
The margins of the few existing streams or pools produce a few scattered willows and
poplars, and the whole country presents a desolate appearance. ‘The plateau is indeed
a continuation of that of Arizona and New Mexico, and though it is depressed in the
valley of the Rio Grande at. El Paso, where the railway crosses, it is still at an altitude
of 3700 feet above the sea. Southward it mounts considerably higher, and on
reaching Zacatecas, the highest point on the railway leading to the Capital, there
is an elevation of 8000 feet. The City of Mexico is situated in a valley surrounded by
ranges of hills clothed towards their summits with pine trees, and reaching an elevation
of about 10,000 feet, while to the south-east are the two lofty volcanoes of Popo-
catepetl and Ixtaccihuatl, the tops of which reach above the snow limit. Numerous
streams descend from the mountains and empty themselves into an alluvial valley with
swampy meadows and form several large lakes, of which Texcoco, Chalco, and
Xochimilco are the most important. It was on Texcoco that the old Aztec Capital
was situated; the houses were built on piles, and the city, which was approached by
a causeway, was destroyed by Cortes at the time of the Spanish conquest in 1519—
1621. The modern town was placed by the conquerors on the swampy ground about
two miles from the eastern margin of the lake, but notwithstanding the altitude it
became one of the most unhealthy capitals in the world, for owing to the frequent
rising of the level of the lake, the town was constantly inundated. Eventually a great
drainage scheme was inaugurated, and for 150 years vast numbers of natives were
employed in cutting a huge dyke for the purpose of draining the overflow of the lake :
but the sodden ground had become so impregnated with sewage that the mortality
was still extremely high, and it was not till 1900 that President Diaz finally completed
the extensive drainage system which now renders the city a comparatively healthy
resort. ‘There can be but little doubt that in former times one large piece of water
covered the whole area of the three lakes, and a map published about 1628 shows
Chalco united with Texcoco, when the surface of the two lakes was far greater
than it is at present. Owing to the drainage system so recently completed, there are

MEXICO. 15:

now several separate lakes draining from one into the other, and finally ending in
Lake Texcoco, which is greatly reduced in size; the water of this lake is brackish,
while that of the other is fresh. .

An account of my journey from Mexico City to Orizaba has been already given
(p. 9), so, after stating that I found myself on a limestone formation surrounded by
rich vegetation, with the high peak of Citlaltepetl towering above, I will refer to
Dr. Gadow for his description of the ascent of this volcano in company with his wife
(‘Through Southern Mexico,’ chap. iii. 1908). He gives in some detail the change
of climate at various altitudes, and its consequent influence on the Flora and Fauna,
which is of special interest here, as conditions somewhat similar prevail on nearly all
high mountains in the tropics.

Starting from Orizaba, Dr. and Mrs. Gadow camped near the village of Xometla at
an altitude of 8600 feet, where they remained for a few days exploring the neighbour-
hood. On their way there they crossed a deep limestone gorge, with fertile vegetation
consisting of a species of P/atanus, magnolias, crotons, and various kinds of oaks,
most of these supporting a luxuriant growth of bromelias, ferns, selaginellas, and
orchids, interspersed with lichens and tillandsias which proved to be * hotbeds
of life.” Northward the open slopes were covered with pasture and clusters of trees
and shrubs, including mimosas, acacias, yuccas, plane-trees, and bamboos. Here was
reached the upper limit of coffee and cotton plantations, while in the damper ravines
tree-ferns were plentiful. Higher up, at the level of the central plateau, maize fields
became scarce, and tree-ferns and datura disappeared. At this altitude a great
change in the vegetation takes place ; there is now but little trace of tropical plants,
and the climate is temperate, moist and fertile, coinciding with the cloud belt. The
vegetation near the camp consisted chiefly of pines, P. montezume and P. liophylla,
with open spaces bordered with deciduous and evergreen oaks, arbutus of two species,
alder and Fuchsia microphylla, with tillandsias in abundance. The larger tillandsias
occur up toa level of 9600 feet, where they suddenly disappear, and mistletoe then
takes their place on the trees. In the ‘barrancas’ or gorges are high trees covered
with creeping aroids and lianas hanging from the branches with abundance of
bamboos and maiden hair fern. Here bird life was almost absent, and only a few tits,
a tree-creeper, a woodpecker, and some blue jays were recorded. A few small
mammals were not uncommon, and the armadillo, which is fairly plentiful in the
lower and tropical country, still exists at an elevation of 8000 feet. Several species of
amphibia and reptiles are also found at this altitude, living chiefly in the bromelias
and other epiphytical vegetation. They have either no lungs or only tiny vestiges
of them, respiration being chiefly carried on through the moist skin. One genus,
Spelerpes, has a wide distribution in Mexico, and S. orizabensis, which leads only
a partially arboreal life, ascends to an altitude of 12,500 feet. Hylodes rhodopis,
which leads the life of a tree-frog, occurs at 10,000 feet, while it also inhabits the

16 PHYSICAL FEATURES ETC.

low hot country of Vera Cruz. Snakes of the genus Crotalus are common in the.
neighbourhood of Orizaba, but disappear entirely in the wet and cloudy zone about
Xometla, being again represented by a small species with a poor rattle at an elevation
of 13,000 feet. Of harmless snakes, only Zropidonotus scalaris. was met with, a
representative of a typically northern genus. A small lizard, Sceloporus microlepidotus,
also has a great vertical range from the hot plains of Oaxaca to the upper tree limit of
Citlaltepetl.

At their highest camp, 12,500 feet, were stunted pines and juniper, while at their
feet grew asphodel and tussock grass, which continued to an altitude of nearly
14,000 feet. At 14,400 feet they reached what appeared to be permanent patches of
snow, but the white summit of the peak, which from this side they found impracticable
of ascent, was still 4000 feet above them. |

At a later date Dr. Gadow, however, ascended the peak itself, an elevation of
18,200 feet, approaching it on the north-east side from La Barbara.

Proceeding in a south-easterly direction from Cordova (2700 feet) there was dense
tropical vegetation on limestone formation all along the foot of the slopes. Eastward
the country showed open savannas, followed by lagoons, swamps, and low forests
in the neighbourhood of Agua Fria. The Rio Papaloapan with its many tributaries
carries an enormous volume of thick yellow water, and for months the low ground
is to a great extent submerged. Further on, Dr. Gadow describes low undulating
erazing land, until he reached the railway which crosses the Isthmus of Tehuantepec.

The eastern slope is clad with humid evergreen forests, but farther west the line
crosses a ridge at an altitude of rather less than 1000 feet, and passes through dense
tropical forests with occasional open patches, but on the Pacific coast a drier type of
vegetation prevails. From Salina Cruz and Tehuantepec north-westwards to the City
of Oaxaca the land again ascends towards the high plateau. The hills at first are
covered with low vegetation, until at 3000 feet pines and oaks appear, but nowhere is
there continuous forest. The general character of the country is dry. Deep gorges
and sandy river beds alternate with an intricate system of hills and patches of
xerophile vegetation. Looking south from an altitude of 5300 feet, the tierra caliente
appears to be densely covered with wood, while to the north is seen a flat, almost
treeless plateau, with here and there outcropping barren ridges of volcanic nature, or
with wooded slopes of Paleozoic formation.

The whole of the Western or Pacific slope is much drier than that of the Atlantic,
and the vegetation is consequently less dense. Pine trees descend to about 2000 feet,
and cacti prevail.

It was on this occasion that, in company with Mr. Julius Flohr, I made a three days’
expedition on mule-back from Mexico City to the Cave of Cacahuimilpa, stopping
at the pretty little town of Cuernavaca en route, from which a fine view of the
western slope is obtained.

MEXICO. 17

The caves, though of considerable size, were on the whole rather disappointing,
and the only animals seen were innumerable bats, which tainted the whole atmosphere ;
as regards other living creatures, even after a diligent search, neither Mr. Flohr nor
myself succeeded in discovering any traces of insects. A Lepisma and a beetle are,
however, reported as having been found by other collectors.

Gadow’s account of the Rio Balsas or Mescala basin informs us that it is bordered
on the south side, parallel with the Pacific coast, by a long high range of mountains,
attaining an altitude of 10,000 feet, densely wooded and intersected by deep gorges,
while the river beds, which form the only available roads, occasionally widen into
meadows.

At Chilpancingo there is a wind swept, shallow depression of cretaceous formation,
surrounded by sparsely wooded hills with meadows on the top of the ridge. ‘To the
west the slopes of the Sierra Madre del Sur are covered with rich forest growth: oaks,
dwarf palms, and pines abound, higher up oaks, pines, and arbutus, and, finally, pines
alone, form dense high forests; while in the gorges, especially within the cloud-belt,
most luxuriant undergrowth prevails. Omilteme (7100 feet) where many specimens
were obtained, is situated in these mountains.

At Cumbre de Los Cajones, a pass of 3500 feet marks the beginning of volcanic
formation. The typical ‘tierra caliente,’ with an essentially tropical flora and fauna, is
found on the southern slope of the main ridge, coinciding with what is officially known
as La Costa. The upper limit may be put at 1000 feet, but the country loses its
tropical character on the ridges, which rise higher than 1500 feet.

From Coquillo to Chacalapan (700 feet) there is tropical life, and from thence to the
coast across lower ridges, the subsoil consists of gneiss and granite in rapid decom-
position, while wooded ground and open pastures are also found. The rivers during
the winter season are frequently dry, but near the granite bound coast are lagoons
mostly containing pure water ; a broad belt of almost impenetrably high forest extends
in places to the sea. Mangrove swamps are permanent features of the landscape, but
in the rainy season many parts of the forests are inundated, and over nearly the whole
of the coastal district a dense mass of tall herbs usurps the place of brushwood in the
forest belts.

Throughout the State of Guerrero large collections, chiefly consisting of birds and
insects, were made for us by Mr. and Mrs. H. H. Smith, who spent a considerable
time in the district, paying special attention to the fauna during the time I employed
them in Mexico. ;

We have no detailed account of the States of Durango, Sinaloa, and Sonora, but the
general aspect is barren and rocky, and although I sent one or two collectors to
those districts, I gained no accurate information as to the physical features.

BIOL. CENTR.-AMER., Introd. Vol., January 1915. D

18 PHYSICAL FEATURES ETC.

REVILLAGIGEDO ISLANDS.

Neither Salvin nor I were able to visit these islands, and the only information
respecting the physical features and distribution of the avifauna is from the
account given by Mr. A. W. Anthony in the ‘ Auk,’ xv. pp. 311-318, after his visit
in 1897.

The Reyillagigedo Islands lie to the 8.W. of Cape San Lucas, Lower California, and
consist of a group of four islands, at some considerable distance apart, but all are of
volcanic origin and in general appearance extremely rough and broken. Socorro, the
largest of the group, is about 240 miles south-west of Cape San Lucas and about 285
miles to the westward of Maria Madre, the largest island in the Tres Marias group,
off San Blas. It is estimated to be about one hundred square miles in extent, and rises
to a height of 4000 ft. in the centre, where there is an extinct voleano. ‘The greater
part of the island is covered with a dense mass of undergrowth which it is almost
impossible to penetrate, especially on the north and north-west, or weather side.
‘Trees are abundant there, but do not exceed forty or fifty feet in height, though
usually covering a considerable area with their spreading branches. On the south
and east the trees are mostly confined to the cafions, where they are smaller than on
the northern slope.

Clarion Island lies approximately about 200 miles westward of Socorro and some-
what further south, and has little in common with the other islands, either in fauna
or flora. Its length is about five miles, and width one mile, the ground rising about
1500 feet above the sea. A few low trees or shrubs, the largest not over ten feet in
height, are scattered along the main plateau, and in a few places extend to the level
ground on the south side, which lies between the Mesa and the coast. Nearly the
whole of this flat ground is covered with a dense growth of cactus (Platopuntia),
over which has grown a mass of vines, and a passage through this belt can only be
made by the diligent use of a bush knife. At a short distance from the beach were
found two small shallow ponds, which contain water during the rainy season only,
but as the high tides evidently overflow the barriers and flood them with sea
water, it is doubtful if they are ever otherwise than brackish. At the time of
Mr. Anthony’s visit, the ponds had been dry for some months, and no water was
found upon the island.

As might be expected from the position and vegetation of Clarion, the birds were
quite different from those on Socorro and San Benedicte. The only land bird common
to any two islands was the Raven, but on Socorro it was not seen.

San Benedicte is a small island about three miles in length with an average width
of half a mile, and lies 35 miles north of Socorro. ‘There is little vegetation, but
wherever sufficient soil is found, there is a heavy crop of coarse grass growing five or
six feet in height, and rendering progress extremely difficult. The barbed seeds

REVILLAGIGEDO AND TRES MARIAS ISLANDS. 19

penetrate a man’s clothing, and were found to be even more unpleasant than the
cactus thickets on Clarion Island.

Roca Partida, or Divided Rock, lies 65 miles north-west of Clarion, and is the
fourth of the group. It is of considerable elevation and has the appearance of a ship
under sail.

The number of species of birds observed by Mr. Anthony are as follows :—Socorro
(24—13 of which were generally distributed land birds), Clarion (17—10 of which
were land birds), San Benedicte (11—including 3 land birds).

TRES MARIAS ISLANDS.

As in the case of the Revillagigedo Group, neither Salvin nor I visited the Tres
Marias Islands; but Alphonse Forrer collected in Maria Madre on our behalf and
obtained a large number of specimens. For a description of the physical aspect of
the islands we ate indebted to Mr. E. W. Nelson, who wrote a ‘General Account
of the Tres Marias Islands with Reports on Mammals and Birds,’ supplemented
by Notes by other authors on the Reptiles, Crustacea, and Plants (North American
Fauna, No. 14, 1899).

The islands are situated about 65 miles off the west coast of Mexico from San
Blas, between lat. 21° and 22° and long. 106-107, and consist of a group comprising
Maria Madre, Maria Magdalena, Maria Cleofa, and San Juanito, arranged in a north-
westerly and south-easterly direction.

About 20 miles from the mainland lies the small island of Isabel, about 1 mile
long and 150 feet high. It chiefly consists of the remains of an old volcano, and
a small crater still occupies the centre. ‘There is evidence of its having once been a
much larger island, though now apparently sinking. ‘The soundings in the channel
between the islands and the mainland gradually deepen to nearly 300 fathoms, but
west of the group the sea bottom falls rapidly to over 1500 fathoms. |

Maria Madre, the largest island, measures 8 by 15 miles, and rises over 2000 feet
above the sea. The interior is occupied by a mountain ridge extending almost the
whole of its length, but descending to a gently sloping area at each end. ‘The eastern
side has the longer slope, while the western or seaward face is much more abrupt,
thus corresponding with the formation of the mountains parallel to the coast on the
adjacent mainland. Both slopes are scored at intervals with cafions, which usually
descend ina nearly direct line to the sea, and along the lower slopes of which, Spanish
cedars and wild figs are grown, certain trees attaining a great size. Generally speaking,
the forest is low and scrubby near the shore, but increases in luxuriance farther up the
slopes. In its primeval condition before the advent of wood cutters, a fine example
of typical forest growth must have been presented here.

North of Maria Madre, and separated by a channel 4 miles wide and 5 or 6 fathoms
D2

20 . PHYSICAL FEATURES ETC.

deep, is an islet 3 or 4 miles in diameter and only about 100 feet high; there is a
narrow border of buffs along the northern shore, thus forming an exception to the
other islands, which are mountainous and rise in successive slopes to the culminating
point near the centre. Here the vegetation largely consists of bush and serubby
trees 8 to 15 feet high, with many agaves on the sandy southern end.

South-east of Maria Madre, and separated by a shallow channel 8 miles wide, is
Maria Magdalena, roughly triangular in outline, and 7 or 8 miles across, rising in the
centre to an altitude of about 1500 feet; south-east again lies Maria Cleofa, the last
of the group. In shape it is irregularly rounded, and about 3 miles in diameter;
the altitude is apparently much less than 1320 feet, as recorded on the charts.
The channel between the two last named islands is about 12 miles wide and much
deeper than the others. Maria Magdalena and Maria Cleofa have a central
mountainous elevation from which cafions descend in all directions to the sea.

The north-eastern points of both islands are low, flat, sandy areas of limited extent,
while the western faces are rocky and precipitous. Permanent fresh water is very
scarce on all the islands.

When visited near the end of the long dry season in May 1897, most of the
herbaceous plants were withered. ‘The general appearance of the vegetation was,
however, the same as in similar situations on the mainland. The most noticeable
plants were Spanish cedar (Cedrela), 3 species of wild fig, 2 of Pithecolobium, 5 of
Solanum, 2 of Ipomea, a Passiflura, Cassias, Euphorbias, a large Agave, a large Cereus,
and 2 Opuntias.

The following is a summary of the species of animals and plants known from the
Tres Marias in 1897, as quoted by Mr. Nelson :—Land mammals, 11 (7 peculiar) ;
birds, 83 (24 peculiar); reptiles, 18 (1 peculiar); freshwater fish, 2; freshwater
shrimp, 1; land molluscs, 6; plants, 136 (12 peculiar). Two species of bats found
by Forrer were not met with by Mr. Nelson, and he was of opinion that both were
stragglers from the mainland.

The relative situation of this group of islands, all with narrow, shallow channels
between them, shows conclusively that at one time they formed a single island at least
45 or 50 miles long, and at a still earlier stage they must have been connected with
the mainland. One of the strongest proofs of this former connection is shown by the
correspondence between the fauna and flora. The breaking down of the original
island into several smaller ones and the continuous encroachment of the sea appear
to indicate that the subsidence is still in progress. The mainland in Tepic near the
coast was within a comparatively recent period the scene of great volcanic activity,
and the Tres Marias Islands bear evidence of having undergone various oscillations,
while the marine deposits of Maria Madre are further indications of the recent
change.

YUCATAN. ‘21

YUCATAN.

During my stay in Mexico in 1887-1888, I determined to visit the province of
Yucatan with the object of exploring the wonderful ruins of Chichen Itza. Taking
my passage in a steamer from Vera Cruz, I landed at Progreso, a port of Yucatan
situated on a spit of sand separated from the mainland by marshy swamps, which,
during the stormy northers that prevail in winter, is occasionally inundated.

After spending a day or two at Merida, the capital, I visited Dr. Gaumer at Izamal
about fourteen leagues distant. ‘This American gentleman, long resident in the
country, had made considerable collections of ‘birds and insects, some of which he
had previously forwarded to us in England.

My original intention had been to ask him to accompany me to Chichen Itza, but,
owing to the disturbed state of the Indians in the vicinity, he advised me not to attempt
the journey, volunteering instead to go with me to Ticul and Uxmal, a journey we
performed on horseback. On leaving Merida we passed through a forest with patches
of open ground, some of which were cultivated with Indian corn and an agave,
from which a fibre called ‘sisal’ is obtained and exported in large quantities. The
name ‘sisal’ is derived from an old port on the north coast, six leagues from Progreso,
from whence the fibre was originally shipped. As we proceeded further south, the
forest trees became larger, but still not of the great size usually found in the tropics.
The ruins of both Uxmal and Ticul have been very much despoiled, a vast number of
the stones having been carried away for building purposes, while many of the carved
pieces formed part of the ‘hacienda’ at which we resided. Froma detailed account
of these ruins when described by Mr. A. P. Maudslay in the ‘Archeology’ of the
‘Biologia,’ it is evident that they were enormously reduced in size since the visit In
1839 of J. L. Stephens, who published in 1843 an account of them in his ‘ Incidents of
Travel in Yucatan,’ with admirable illustrations by Catherwood.

The peninsula of Yucatan is flat and of a recent limestone formation ; there is a low
range of hills which stretches from a point a few miles south of Merida to the
neighbourhood of Peto some distance south of ‘Ticul, but nowhere exceeding 500 feet
in height.

The coast is very low and swampy, while further inland are forests, which in a
few cases have been cleared, but the whole country is very sparsely inhabited. ‘The
southern part is, so far as I could learn, but little known, but it is said to be largely
covered with forest and the trees are much finer than those in the north.

The following description of the country is mainly taken from Dr. Gaumer’s notes
published in Boucard’s account of a ‘ Collection of Birds from Yucatan’ (P.Z. 8. 1888,
pp. 484-439), supplemented by my own observations in 1887-1888. At Tizimin the
country, like the rest of Northern Yucatan, is on a low level, but to the north east and
south lie vast forests, for the most part uninhabited since the migration of the Indians

22 PHYSICAL FEATURES ETC.

half a century ago. This region is filled with ruins both ancient and modern, but of
the former very few of any size remain. Some of the ranchos have, however, been
re-peopled and most of the birds have been obtained in the vicinity of the clearings.
At Yok Jonat Ku there is a large forest where the trees are high and the ground
comparatively open; here the magnificent turkey Meleagris ocellata is still to be found.
At one time this bird was distributed all over the peninsula, but owing to the depre-
dations of the Indians, who esteem it highly as an article of food, it is now almost
exterminated.

Lagartos is a sea-port town at the mouth of the river, or more properly an arm of
the sea bearing the same name, and innumerable streams or—as Dr. Gaumer believed—
subterranean rivers find an outlet there. The waters are very salt, and in the dry
season are even more saline than the sea itself. ‘This so-called river is broad and
shallow, bordered by a dense growth of low bush, behind which he marshes of salt or
brackish water, and here in June and July thousands of flamingoes in their finest
plumage were seen by Dr. Gaumer, while swarms of other sea-birds were always in sight.
The innumerable hosts of mosquitoes which come with the first rain impeded the
work here, and the intermittent and pernicious fevers render collecting both difficult
and dangerous. The country generally has no surface water, and the only supply is
from the Aguadas and Cenotes (Senotes or Jonats), as they are called by the natives.
Fortunately the Aguadas, which are said to be of natural formation, but which appear
to have been reconstructed by the ancient Indian races, are very numerous; they
consist of a deep excavation in the earth, sometimes circular in form, but giving the
idea of having been at one time quadrilateral, and from fifty to one hundred feet in
diameter. They contain water all the year round, though never of any great depth.
‘The sides being inclined, they form natural drinking places and are much frequented
by animals and birds—so much so, that the collector usually obtains a good number of
specimens in the vicinity. .

The Cenotes are probably natural openings in the earth with steep walls of limestone
frequently sixty feet high; they vary in size and shape, but always contain clear, fresh
water. They are believed to be openings to underground rivers, and are frequently
found in immense caves with a narrow circular mouth ; at the water's edge there is no
resting-place and no approach except by the steep sides. Vultures, owls, and similar
birds nest in the walls. The caves are also frequented by swallows, bats, and motmots,
and reptiles are said to occur in immense numbers. ‘The water contains numerous
fish belonging to the Siluride, and in the shallow open water-holes near the coast
there is said to be another species belonging to the same group, but Dr. Gaumer
was, unfortunately, unable to capture a specimen. ‘The distribution of these Siluride
confirms the belief that underground rivers in Yucatan do exist.

A very interesting description of the climate (in 1878-1879) is given by Dr. Gaumer *,

* See Boucard, P. Z.8. 1883, pp. 434-462.

YUCATAN. 93

who first reached the country in the middle of October 1878. The summer rains had
ceased about ten days previously and the weather had been good, but throughout
Qctober, November, and December ‘norther’ followed ‘norther’ every ten to
fourteen days, accompanied by light drizzling rain which lasted generally from two
to four days with increasing cold. In January there were four moderately heavy
rainfalls with strong ‘northers’ and cold nights. In February there were five
‘northers’ and one with very heavy rainfall accompanied by hailstones of such size
that they were quickly gathered up and by many people placed in bottles thinking that
they might so be preserved. From February 26th to May 23rd no rain fell and the
sky was cloudless for weeks at a time. The heat during the day gradually increased,
until in April and May it was almost intolerable, and on account of the dry air and
clear sky the radiation was so great that the nights were disagreeably cool, though
generally so balmy and pleasant. ‘The birds disappeared as the dry season advanced,
when only a few common resident species round the ranchos and Aguadas were to be
found. On May 23rd the first summer rains commenced and were followed by daily
showers at mid-day. All nature changed as if by magic, new leaves grew, and the
forests were again populated with songsters. In June the rains began at 11 a.m. and
ceased at 2 P.M. with an almost daily regularity. In July they began at 10 a.m.
and ceased at 3 or 4 P.M., but never earlier. In August it rained from 10 a.m. till
nightfall and sometimes later. During these last three months there were from five
to eight days in each month upon which no rain fell. The heat was almost insup-
portable, even to the natives, and yellow fever raged in the towns of the interior. In
September the rains lasted from 8 or 9 a.m. till midnight and not infrequently all
night, but the weather became milder. Reptiles were almost the only things to be
found. In October 1879, during the first twenty-seven days, rain fell in torrents and
almost incessantly, the sun was seen but on four or five days and the stars appeared in
patches on five nights only, and not five consecutive hours were fine during those
twenty-seven days. Yellow fever gave place to bilious fever. Insects were rarely seen,
birds almost entirely disappeared, and any skins were worthless, as either owing to the
heavy rain or to some other cause the feathers had not yet commenced to grow.

An account of Yucatan would scarcely be complete without some particulars of the
distribution of species and a comparison with that of the islands off the coast and in
the Bay of Honduras, which were visited on our behalf by Dr.Gaumer. In a summary
of the island-birds examined in detail by Salvin (‘ Ibis,’ 1890, pp. 84-95) he arrives at
the conclusion that they split up naturally into three groups. Leaving out Meco, the
exact position of which is uncertain, he considers that those from Holbox and Mugeres
may be classed together, Cozumel by itself, and Ruatan and Bonaca (Guanaja) by .
themselves, though all show a strong affinity to the birds of the mainland. The two
last named islands are stated to be very different in their physical features to the
others mentioned ; they are of high altitude and attain an elevation of 1200 feet, and

24 PHYSICAL FEATURES ETC.

have the upper parts covered with pines, in contrast to the low ground and recent
coral-limestone formation of the remainder.

The total number of species obtained was 214, of which 79 are migrants from North
America, and of the remaining 135 species, 27 are birds which frequent the sea coast.
The table of distribution of the 108 non-migrating species shows that the affinities of
the birds of these islands as a whole are largely on the side of those on the mainland,
the West Indian element being very slight. Northern Yucatan and the islands
adjoining are separated from Cuba by a depth of over 1000 fathoms, and the Bay
Islands from Jamaica by over 500 fathoms. Had there been any recent land con-
nection, a supposition which the similarity of the birds alone would justify, the main-
land as well as the West Indies would hardly fail to show such a connection in a much
more pronounced manner, and we should not find the strong contrast which exists
between the faunas of Cuba, Jamaica, and the mainland, but a larger number of features
in common ; this contrast is still more marked in the Lepidoptera Rhopalocera.

The alternative supposition to account for the West Indian element in these islands
is that the birds have reached them at no distant date by flight. When we consider
that the trade wind blows in the direction of Yucatan and this coast for several months
in the year, it is scarcely a matter of wonder that some West Indian birds stray so far
west. Cozumel appears to have been separated from the mainland for a considerable
period, during which time it has received casual immigrants from the West Indies,
from North America, and from the mainland, some at a distance of time sufficient to
allow of their modification. There are 159 species of birds, 65 of which are migrants
and 27 of very wide range.

My stay in Yucatan was very short, and owing to the state of my health, which
rendered me unfit for much exertion in a country so little explored, I was able to do
very little collecting. ‘The specimens acquired, however, were, as before, mostly due
to the enterprise of Dr. Gaumer, who employed natives to collect, but they were
necessarily from a limited area. My trip ended very much as it began, for I was
obliged to return to Progreso, and from thence by steamer to Vera Cruz.

BRITISH HONDURAS.

British Honduras, or the colony of Belize, as it is often called, situated on the south-
eastern shore of the peninsula of Yucatan, is about 160 miles from north to south and
60 at the widest part. The navigation of the coast is both difficult and dangerous,
on account of the numerous cays and coral reefs with which it is bordered. In the
neighbourhood of the town of Belize, and for some distance inland, the ground is low
and swampy and thickly clothed with mangroves and tropical jungle. Further west
there is a narrow belt of alluvial soil, beyond which, and parallel to the coast, are
tracts of arid sandy land called ‘pine ridges,’ from the pine trees with which they

BRITISH HONDURAS. 25

are covered. Still further inland are the ‘ Cahoon Ridges,’ clothed with palm trees,
while beyond are broad savannas studded with clumps of trees and intersected with
streams. The Manatee Hills rise in a further succession of ridges parallel to the
coast, and are from 800 to 1000 feet in height, while to the south the Coxcomb
Mountains attain an altitude of 4000 feet; further inland there are said to be a
succession of valleys and hills at altitudes varying from 1200 to over 3000 feet above
sea-level, but this part is very imperfectly known. The climate near the coast is
generally hot and damp, but tempered by the trade winds, and though the annual
rainfall is said to be about 100 inches, the country is tolerably healthy. Unlike the
rest of Central America, British Honduras is not subject to earthquakes ; it appears
to be entirely outside the volcanic area, which otherwise extends from Mexico to
Western South America.

In 1862 Salvin, as stated on p. 5, went from Coban by way of Peten, down the
Belize River to the town of Belize on the coast, but unfortunately he left no details of
the country through which he passed. His intention was to proceed direct to Yzabal,
and thence back to the interior of Guatemala, but finding no vessel ready to sail, he
hired a schooner and occupied the time in exploring some of the numerous atolls and
coral-reefs which line the coast, and later published an interesting account of this
expedition in ‘The Ibis’ for 1864. He described the Barrier Reef as extending
from Ambergis Cay to Ranguana Cay, its most northerly point; this last cay is twenty-
five miles from the coast, so that the reef, instead of running more or less parallel with
it, forms an angle enclosing a long lagoon, which, as weil as the reef, is studded with
numerous cays. Nearly due east of the town of Belize, outside the Barrier Reef, and
separated from it by a deep channel, lies the Atoll of Turneff, within which several
lagoons are included. Fifteen miles eastward of Turneff lies another atoll, called
Lighthouse Reef, on the eastern margin of which are four cays—Long Cay, Middle
Cay, South-West Cay, and South-West-of-All Cay; the remainder of the reef consists
of a line of breakers, showing here and there a stranded log or a protruding spit of
sand. It will be easily understood that these reefs, many of which are covered with
mangroves and coco-nut palms, form an ideal place for sea-birds, and as Salvin’s visit
took place at the height of the breeding season, he procured a large number which he
had not previously obtained.

In addition to these sea-birds, Salvin mentions two humming-birds, two tyrants,
a warbler, a mocking-bird, an osprey, an ibis, egrets, etc. ‘The paper quoted is too
long to reprint in detail, but it is still the only account known to me describing
the cays in question. His subsequent visit to the lagoons on the Pacific Coast of
Guatemala in 1863 is referred to under the heading for that country.

BIOL. CENTR.-AMER., Introd. Vol., January 1918. E

26 . PHYSICAL FEATURES ETC.

HONDURAS.

The Republic of Honduras is bounded by the Bay of that name and the Caribbean
Sea on the north, by Guatemala on the west, and Salvador, the Pacific Ocean, and
Nicaragua on the south; it includes the islands of Ruatan, Bonaca (Guanaja), and
the islands adjacent. The general aspect of the country is mountainous, and it is
traversed by ranges and hills radiating from the base of the Cordillera. ‘The main
chain, which does not approach within 90 or 60 miles of the Pacific, is not an
unbroken one, as it turns back and forms basins or valleys, within which are collected
the head-waters of the streams which flow in the direction of the Atlantic Ocean.
Viewed from the Pacific, the mountains present the appearance of a great natural
wall, with a lower range bristling with volcanic peaks between it and the Western
Ocean. The Cordillera proper forms an irregular line from north-west to south-east,
interrupted, however, by the great transverse depression of Comayagua, which extends
about 40 miles to the north with a width of from 5 to 15 miles, and contains the
Humuya River, which discharges its waters into the Atlantic; while to the south it
forms the valley of the Goascoran River, which flows into the Pacific. The whole
country has a great diversity of surface and elevation, with fertile valleys and high
plains, affording every variety of climate.

Some notes on the aspect of this country were published in ‘ The Ibis’ for 1860 by
G. Cavendish Taylor. He crossed from the Pacific to the Atlantic Coast, and many
of the places described have been mentioned in the ‘Aves’ section of this work.
Arriving in December 1857, from Panama, at La Union, Salvador, he crossed to the island
of Tigre, and thence to La Brea in the Gulf of Fonseca, which is studded with densely
wooded volcanic islands. Tigre abounds with scorpions and large hairy spiders, and
many birds were obtained near an old crater, now a lagoon full of reeds and floating
grass. Mr. Taylor visited Aremecina, Caridad, San Juan, and Lamani, and so over the
tolerably level plain mostly covered with forests to Comayagua. In the vicinity of
the town were cactus bushes on one side, dense jungle intersected by rivers and woods
on the other, and high mountains bounding the plain. Continuing his journey towards
the Atlantic, Siquatepeque was reached by a route crossing the top of the mountains
at an elevation of 5000 feet—here again was open plain, 2600 feet above sea-level,
and surrounded by mountains; then after passing over undulating ground covered
with pine trees, giving it a park like appearance, he arrived at Taulevi. Here
arrangements had to be made for the journey through the dense forests to the Lake
of Yojoa, some three leagues distant. Men had to be sent forward to clear a path
and engage boats for the passage down the lake. The route lay at first by open

HONDURAS.—GUATEMALA. 27

savannas and wooded hollows, but soon a broad, rapid stream was crossed, and entrance
was made into the dense forest, through which a road had to be cut, but further on
the trees were lofty with huge buttresses at the base, and the undergrowth was not
very thick. It was nearly dark when the river, flowing from the lake, was reached
and the canoes launched. The current was slight, the water deep and still, and the
banks were covered with high trees and dense forests, every bush and bough was alive
with fire-flies, and the cries of night-hawks, coupled with the croaking of innumerable
frogs, made no inconsiderable noise. The wind was ahead and occasionally so strong
that the travellers were unable to proceed until it lulled, but at dawn they had
completed twelve miles and had reached their destination. ‘Two days were spent at
Agua Azul, so called from the colour of the deep spring which rises near the ‘hacienda’
and flows into the lake. Numerous interesting birds were seen among the reeds and
alligators (Crocodilus americanus) were not uncommon, while every tree and blade of
grass swarmed with ‘garrapatas.’ Leaving the lake, which was surrounded by high
mountains, the journey was made principally over savannas and open ground to the
town of Yojoa, and vid Potrerillos to the Atlantic. In the forest the route lay for
miles through vistas of palm trees and bamboos, which shaded the path with their
feathery branches, but unfortunately prevented the deep mud-holes from drying up.
After leaving San Pedro, where brown monkeys with white faces were seen, a high
range of mountains was crossed and Omoa reached on February 14th.

G. C. Taylor enumerates one hundred species of birds, and G. M. Whitely sub-
sequently sent us a collection of 135 species from the same country. These were
named by Salvin and a list of them published in the ‘ Proceedings of the Zoological
Society’ for 1870, pp, 835-839. Whitely’s skins (520) were obtained in 1869 in
the vicinity of Puerto Caballo (Cortes) *, Julian, Medina, and San Pedro. The three
last named places are situated in the low forest-lands on the Chamelicon River; San
Pedro, the farthest inland, is not more than 30 miles from Puerto Caballo, now the
Atlantic terminus of an incompleted inter-oceanic railway.

After examining the collection carefully, it became apparent that the Ornithology
of this part of Honduras scarcely differs from that of the thoroughly explored iowlands
of Vera Paz.

GUATEMALA.

Guatemala is coterminous on the north and west with Mexico, the flat low lying
peninsula of Yucatan extends to the north-east, British Honduras, the Caribbean
Sea, and the Republics of Honduras and Salvador are on the east and south-east,
while on the south-west lies the Pacific Ocean. The greater part of the country is

* Often confused with Puerto Cabello in Venezuela.

E2

28 PHYSICAL FEATURES ETC.

mountainous, the main chain of the Cordillera forms the watershed, and at a mean
elevation of 7000 feet runs nearly parallel to the Pacific coast at a distance of about
fifty miles from it. The steep slope on the Pacific side is broken by many voleanoes,
while towards the Atlantic the land sinks in a gentle incline with subsidiary ranges
extending nearly to the water’s edge. Of the voleanoes several are active, the most
noted is the Fuego (14,070 feet) with its twin sister the Agua, so called because
in 1541 a lake, which occupied the centre of the crater, was discharged by a great
eruption on to the former capital below and the city now called Ciudad Vieja was com-
pletely destroyed by water. Both volcanoes are clothed with dense forests from about
7500 feet to 10,000 feet, above which level there are scattered pines for 1000 feet
or more, of which stunted examples are even to be found in the extinct crater of the
Agua. Below 7000 feet the forest has been cleared for cultivation, and only parts
are now clothed with a dense growth of scrub. ‘The chief rivers are the Usumacinta,
which flows into the Gulf of Mexico, and the Motagua and Polochie, which fall into
the Bay of Honduras. Those flowing into the Pacific Ocean are short and rapid, as
the fall from the Cordillera is very steep.

The so-called ‘coast country,’ however, extends a long way inland, as during the
wet season the torrents which descend from the Cordillera are charged with volcanic
sand and disintegrated scorie, and when discharged into the ocean they are cast
back by the waves and the deposit forms a line of sand-bank. The constant heaping
up of this bank often closes the mouths of the smaller streams during the dry season,
and when the current is not sufficient to reduce the sand-bar the water expands inside
the beach, forming lagoons and marshes along the whole coast. ‘These lagoons are
a favourite resort for waders and sea-birds, of which Salvin later obtained a large
number, as well as a considerable quantity of fish.

As previously mentioned, Salvin made no less than four expeditions to Guatemala,
but I will only describe the physical aspect of the country we travelled over together,
alluding, however, to those parts which he visited alone and giving extracts from
some of his scattered papers published in ‘The lbis’ and other magazines. We
landed in September 1861 at Yzabal, on the Golfo Dulce, and after a short stay we
proceeded towards the Capital. The neighbourhood of Lake Yzabal is covered with
dense forest extending beyond the Mico Mountains, which we crossed into the
Motagua Valley in order to reach Quirigua. Here we spent a few days, and then
proceeded up the river valley to Zacapa by mule path, the country gradually becoming
drier and the vegetation more arid; cacti and thorny shrubs became abundant, taking
the place of more luxuriant plants. Indian settlements were found at intervals of
every few miles, where the brushwood had been cleared for the cultivation of maize
and coffee-trees, which were growing in small patches. Much the same character of
dry country prevailed throughout the journey to Guatemala City, which is situated on

GUATEMALA. 29

the Cordillera at an altitude of about 4500 feet above the sea. From the Capital we
visited Duefias, a village on the Pacific slope situated in a plain at the foot of the
Volcan de Fuego and between it and the Volcan de Agua. Here we spent about
three months, exploring the country and making frequent excursions into the forest
of the Volcan Fuego, which furnished a great contrast to that near the coast; the
high trees were the only corresponding feature, but both vegetation and climate were
entirely different. Instead of the incessant noise of the buzzing of myriads of insects,
life seemed almost extinct and a dead silence reigned throughout, broken only by an
occasional gust of wind or the fall of some rotten tree. The mountain itself is
furrowed with deep ravines, called ‘barrancas,’ the sides of which are exceedingly steep
and quite impassable, and in ascending the mountain, care must be taken to keep on
the top of the ridges between them. ‘The forest shuts out the view of the surrounding
country, consequently landmarks are not available, and as one ravine almost exactly
resembles another it is an extremely easy matter to lose one’s way by inadvertently
following the edge of a new ‘barranca.’ ‘The lower part of the forest up to about
7000 feet has been cleared for cultivation, but quickly reverts to a dense growth of
scrub, above which is found a belt of evergreen-oaks followed by deciduous trees
of various species, amongst which the remarkable Cheirostemon platanoides mingles
in the highest range with alders. ‘Then follows more open ground with pine trees
and coarse grass, but the trees become stunted as the ascent increases, and finally
disappear at an elevation of about 11,000 feet. From thence to the summit of the
Fuego the cone is composed of cinders and ashes interspersed with short coarse grass.
The mountain is divided into two peaks or cones near the summit, connected by
a narrow ridge of cinders, the southern and higher peak is still active, and from it a
perpetual column of thin smoke is always plainly visible. ‘This cone is very steep, and
the climb to the edge of the crater itself is exceedingly laborious, as the foot sinks at
each step deep into the ashes. The view, however, from the point well repays the
trouble of the ascent. Southward the eye travels a distance of 50 miles to the coast,
far beyond again is seen in dim outline the horizon of the Pacific Ocean, while
below on the other side lies the deep abyss of the crater itself. The northern cone
is more or less covered with coarse grass extending to the summit, while the interior
of the crater has been almost filled with the eruptions of the more recent southern
volcano; but signs of internal fires are not wanting, as jets of steam and sulphurous
vapour are still seen issuing from the fissures in the rocks. In one of the hotter
crevices I found a vigorous plant of Lycopodium clavatum and a Selaginella taking
advantage of the warmth and moisture and growing with wonderful luxuriance at an
altitude of nearly 14,000 feet. The descent was by no means easy, as there was no
track to mark our way, but we had fortunately taken the precaution of slashing
the trees with our big knives or ‘machetes’ on our way up, which indicated our

30 PHYSICAL FEATURES ETC.

route sufficieatly to enable us to return to camp in safety. ‘The Volcan de Agua is
very similar to the Fuego, though somewhat less in height, but there is a fairly good
mule track nearly to the summit. This track is frequented by the Indians, who
ascend the mountain for the purpose of charcoal-burning and also in search of ice,
which they found in the old crater in sufficient quantity to supply the needs of the
Capital at that time.

Having made considerable collections, we returned to Guatemala City and, recrossing
the Motagua River and the Chuacus Range of mountains, took up our abode at San
Gerénimo, a sugar-cane plantation in the plain of Salama, in Baja Vera Paz. The
surrounding mountains are clothed with forest composed of various trees, including
pines. The plain itself is arid, except when irrigated for cultivation, as at the
Hacienda of San Gerénimo. We next proceeded to Coban through the district north
of the plain of Salama. The road soon leaves the plain, and the broken country is
covered with scrub and forest, the rainfall being much greater as one approaches Coban
than on the Pacific side of the Cordillera. During the rainy season there is usually
a severe thunderstorm in the afternoon, followed by a clear sky, but during the dry
season little or no rain falls and vegetation suffers greatly. In Alta Vera Paz and
towards the Atlantic rain apparently falls at all seasons and all hours, and vegetation
is consequently much more abundant. :

At Tactic, a forest district near the head of the Polochic River, our porters failed
to arrive, and we were forced to spend the night without our baggage. It was so
bitterly cold that in the morning the ground and even the backs of our mules were
covered with hoar-frost. A few days later, on our return journey, the effects of the
unprecedently low temperature were plainly visible on the vegetation around. On
reaching Coban we found a large Indian village where the inhabitants were born
collectors, and very soon they brought in, in almost embarassing numbers, specimens of
birds, frogs, toads, lizards, snakes, and insects of all kinds. ‘The natives there were
specially expert in the use of the blow-pipe, with which they killed most of the
smaller birds. The weapon consisted of a straight piece of hollow wood about
eight feet long, and the projectile, a hardened pellet of clay, fitted closely into the
groove of the pipe and was blown from the mouth by the marksman. In this way
a large number of birds was obtained with little or no damage to the plumage. Such
was the accuracy of aim that, even at a distance of from 15 to 20 yards, many
humming-birds were killed.

After some weeks spent in collecting at Coban we visited Cubilguitz and Choctum
in the low damp forest of Alta Vera Paz, thence travelling towards Salinas in the
humid valley of the Chixoy or Rio de la Pasion, a tributary of the Usumacinta River.
The roads or tracks made by the natives were extremely bad in this locality, the
eround very broken, and the soil a stiff clay, so slippery in places that it was scarcely

GUATEMALA. aL

possible for animals to keep their footing. As there were no villages whence food
could be obtained in this little known district, it was necessary to take a three weeks’
supply from Coban and also to engage a number of Indians to act as porters. It
was somewhat difficult to estimate the amount of food required per person, and for
this purpose we decided to make a preliminary or trial trip extending over three
days. We found that an Indian consumed daily about half his straw hat full of
‘topopoxti’ or baked Indian corn cake, and this with a few onions and ‘frijoles’ or
black beans supplied the necessities of life. Having arrived at the quantity required,
we made up a sufficient number of loads and these were carried by the porters
on their backs.

En route we occasionally discovered a small Indian settlement, where our “‘ mozos”
found shelter in a hut formed of poles and thatched on the top and on two sides.
These were resting places used by the natives on their way to Salinas in search of
salt. Salvin and I preferred, however, sleeping in our hammocks slung to the trees
in the adjoining forest, and as we were each provided with a waterproof sheet, we
slept in dry beds notwithstanding the constant wet nights.

The days were usually fine and were mostly spent in exploring the forest and
collecting birds, insects, and plants. We remained a little over three weeks, till our
supplies were exhausted, and then returned with our spoils to Coban.

Owing to my having contracted an attack of fever and ague in the low ground
at Salinas I was unable to accompany Salvin on his journey to Peten and Belize.
On his second expedition to Guatemala, Salvin had already visited Lanquin and
Cahabon, about three or four days’ journey from Coban. He describes (‘ Ibis,’
1861, pp. 138-149) the country as very wet and covered with forest, the roads
—or rather tracks—impassable for animals, and all baggage had to be carried by
Indians. The forests on the slopes of the limestone mountains were the home of
the Quezal, the royal bird of the Aztecs, as well as of many other birds not found
on the Pacific side, such as members of the families Cracide, Tinamide, etc. Salvin
says: ‘“‘These forests are perhaps more worth seeing than anything in Guatemala,
quite different to those on the West Coast, where the heat is excessive and mosquitoes
and other insects abound and destroy one’s comfort. In these forests it is otherwise ;
no ‘ garrapatas,’ no mosquitoes, and a climate in the dry season which might challenge
any in the world. Most parts are free froma brushwood, and one may ramble where
one pleases without being stopped by dense thicket. What strikes the eye most is
the number of ferns, not only of plants, but of species; every tree is clasped, every
stone clothed with them, besides many of terrestrial habit.”

As soon as I had recovered from the effects of fever, I left Coban for Buenaventura
on the Motagua River in order to collect fish, and the methods employed have already
been described on a preceding page. |

32 PHYSICAL FEATURES ETC.

Salvin then revisited his old quarters at San Gerénimo, and taking his friend Robert
Owen, the proprietor of the Hacienda, with him, he rode over the high land round
Quiché and Totonicapam at an altitude cf 10,000 feet. Here the climate is
temperate, potatoes and wheat are largely grown, and on the uncultivated ground
oaks, pines, and alders abound. Thence, crossing the Cordillera, he proceeded to
Quezaltenango, a large town in the ‘ Altos,’ and the capital of a considerable district,
which he describes (‘Ibis,’ 1865, p. 187) as a corn growing and sheep producing
highland ; thence to Retalhuleu and on to the port of San José. At Retalhuleu he
he «d such glowing accounts of the prospect of obtaining a valuable collection of
sea-birds and fish from the lagoons on the coast that he took a passage in a trading
barque which was going from San José to Champerico to take ina cargo of coffee and
sugar, and succeeded in procuring a large number of specimens.

When Salvin had finished collecting on the lagoons, he made an expedition to a
belt of tropical forest parallel to the coast, but about twelve miles distant. Here it
was that he specially remarked the contrast between the birds of the Pacific and
Atlantic coasts—many of the most familiar birds of the low forest of Vera Paz, the
Tinamide, Columba nigrirostris, and Ostinops montezuma, being entirely absent, nor
does one find the genera Rhamphocelus or Calliste, or the beautiful Cotinga amabilis.
Much of the forest consists of bamboo, with here and there a huge tree standing high
above it. Between this forest and the coast the soil is comparatively unproductive,
bearing the stamp of land reclaimed from the ocean at no very distant date. The
long line of volcanoes suggests a recent upheaval, and the constant discharge of sand
by every river would tend to advance the coast by slow degrees. This low country is
very subject to malarial fever—although Salvin escaped, his two attendants contracted
it. Salvin returned to England soon after this, early in 1863, but ten years later, in
the autumn of 1873, he paid his fourth and last visit to Guatemala, this time in
company with his wife; although he added considerably to our collections the route
taken was much the same as on previous expeditions.

In this description of Guatemala, it must be remembered that when I visited it over
fifty years ago there were no railways. ‘There is now a railway from Puerto Barrios
on the Atlantic Coast up the Motagua Valley to the capital and thence to the Pacific
coast at San José, with a branch running from Mixtan near Escuintla to Retalhuleu
and Champerico. The country, therefore, is at the present time readily accessible by
steamer from Belize and thence from the Atlantic port by train.

SALVADOR.—NICARAGUA. 33

SALVADOR.

Salvador, though the smallest of the Central American Republics, is one of the
most densely populated and largely cultivated, and consequently there is but little
forest. It is bounded by Guatemala on the west and Honduras on the north and east.
The country averages about 60 miles in breadth, and the coast line on the Pacific is
160 miles in length. The seaboard consists of a comparatively narrow alluvial plain,
beyond which is a plateau with a mean elevation of 2000 feet, broken by a number of
volcanic cones lying to the south of the main Cordillera, and the whole Republic is’
very subject to earthquakes and volcanic outbreaks. ‘The general aspect of Salvador
led us to believe that the fauna and flora would be very similar to that of Guatemala
and Honduras, consequently we neither visited the country ourselves nor did we
employ any collectors there.

NICARAGUA.

The Republic of Nicaragua, wedged in between Honduras on the north and Costa
Rica on the south, has a coast-line of about 280 miles on the Caribbean Sea and about
200 on the Pacific. The land gradually decreases in width from north to south, while
the main watershed extends eastward from within a few miles of the Pacific Ocean.
Greytown (San Juan del Norte), at the mouth of the San Juan River, formerly
possessed a fine harbour, but of late years the Colorado branch of the river, which
bifurcates about twenty miles from the coast, now takes most of the water and the old
channel and harbour have silted up. The main geographical feature of the country is
the remarkable depression stretching for about 200 miles from the north-west to the
south-east, parallel with the Pacific Coast and to the central plateau. This depression,
which lies at a mean elevation of about 100 feet, is flooded by two great lakes,
Managua and Nicaragua, which collect the drainage-water of the western provinces and
also that from the eastern range of mountains, finally discharging it through the San
Juan River into the Caribbean Sea, a distance of 120 miles. ‘The Lake of Managua is
about 50 miles in length and 25 in breadth; the level is 16 feet higher than that of
Lake Nicaragua, but the natural outlet, except in high flood, carries but little water,
the surplus passing off by evaporation. The Lake of Nicaragua is about 100 feet above
the sea-level and 150 miles long. Throughout its entire length this great depression
is traversed by a remarkable chain of isolated volcanic cones, which, north of the lakes,
takes the name of Marabios, terminating at the extreme north-west with Coseguina
(4000 feet), and in the extreme south-east in the low wooded archipelago of Solenti-
name and Chichicaste, near the entrance to the San Juan River. These volcanoes
range from 4000 to over 6000 feet, while Momotombo, the highest point in the Republic
of Nicaragua between the Gulf of Fonseca aud Lake Managua, reaches an altitude of

BIOL. CENTR.-AMER., Introd. Vol., January 1915. EF

34 PHYSICAL FEATURES ETC.

7000 feet. To the above mentioned series of volcanoes also belong those on the islands
of Zapatero and Ometepe, in the lake of Nicaragua. The latter, after a long repose,
burst into renewed activity in 1883, and for seven days continued to spread devas-
tation, destroying the crops and compelling the people to take refuge on the mainland.
Several other volcanoes are still more or less active, and in 1835 Coseguina was the
scene of one of the most tremendous eruptions on record. The outbreak lasted four
days, during which time sand and ashes were carried to such a distance that they fell
in Jamaica, Mexico, and Bogota. No rivers of any size flow westward to the Pacific,
but the Lake of Nicaragua receives, near its outlet, the important Rio Frio from Costa
Rica, which, at certain seasons, brings down a vast amount of water. Little is known
of the region of rugged plateaux and savannas occupying fully half the country between
the lacustrine depression and the Caribbean Sea. A large portion of the low ground
is said to be covered with dense forest intersected by innumerable streams, all flowing
eastward to the Mosquito Coast, which is low, swampy, and very unhealthy.

Mr. Thomas Belt, a mining engineer and a well known naturalist, spent over four
years at the gold mines of Santo Domingo in Nicaragua, and published an excellent
account of his travels on his return in 1874. A considerable part of his book is
occupied with extremely interesting observations on the Indians and the natural history
of the district through which he passed, and I am indebted to him for the following
details. Landing at Greytown he proceeded in an open boat up the San Juan River,
which he describes as having a dangerous bar, over which he had to pass; he then
entered a wide channel with shallow water and beds of high grass on one side and a
sandy shore on the other, in which alligators floated about like logs of wood and flocks of
wading birds were seen in the marshes beyond. Proceeding up the river in still water,
he emerged into a wider channel with a stronger current. The banks of the river
were at first low and marshy, intersected by numerous streams fringed chiefly with
palms and beds of wild cane and grass; further up the banks became higher and
drier, and plantations of bananas and plantains were noticed in the clearings of the
forest. About twenty miles above Greytown Mr. Belt reached the Colorado branch
of the river, which now takes the greater part of the water from the lake to the sea
by another outlet. ‘There the banks were hidden by high trees laden with creeping
and twining plants, many of which bore beautiful flowers, while beneath were tree-
ferns with their light green foliage and siender stems. Higher up he passed the mouth
of the Chiripo River, which rises in the interior of Costa Rica and joins the San Juan
about thirty miles above Greytown, It is navigable for about twenty miles from this
point, after which it becomes a rough mountain-torrent, and a mule track leads thence
to San José, the capital of Costa Rica. At Castillo, on the river bank, a considerable
quantity of ‘Ulli’ (ule) or rubber is collected by the Indians, which forms an
important object of trade. This is obtained from a species of wild fig (Castilloa
elastica), a plant with large leaves, differing entirely from that found on the banks of

NICARAGUA. 35

the Amazon (Siphonia elastica), and which is prepared in another manner; the former
is abundant in the forests of Nicaragua and Honduras. ‘The San Juan River continues
with much the same high forest on its banks as far as San Carlos, at the entrance to
the Lake of Nicaragua; it is about 120 miles long and the lake is 107 feet above the
sea, so the water falls a little less than one foot a mile. The height of the lowest pass
between the lake and the Pacific Ocean is said to be only 26 feet, and consequently
this is the greatest depression in Central America between the Atlantic and Pacific.
Owing to the enormous reservoir of water in the lakes, it has frequently been
suggested as a practicable route for a ship canal between the two Oceans.

On reaching the lake a sail was hoisted on board the little boat, and in a couple of
days Belt arrived on the northern shore at Ubaldo, the landing-place for the machinery
and goods destined for the mines at Santo Domingo. Leaving Ubaldo the road
crosses some low rocky hills with scanty vegetation, consisting of spiny cacti, leathery
leaved trees, thorny palms, prickly acacias, and bromelias with sharply serrated
leaves; this being the dry season, the mule track was parched and dusty, though
during the rains it becomes a slough of mud and water. The road led through the
town of Acoyapo, which is in a grazing district with large cattle ‘ haciendas.’ Soon
after this, Belt crossed the range which divides the forest region extending from the
mountains of Segovia to the Caribbean Sea, and separating it from the great lake
depression. The savannas on this side were more humid and the moisture increased
as he proceeded across the upper waters of the Mico River, which enters the sea
at Blewfields. The black margin of a great forest, which had been visible for some
time, was reached in the neighbourhood of Santo Domingo; the ranges of irregular
hills running mainly east and west were covered with vegetation, which was usually
enveloped in a dense mist and produced a most depressing effect. The last part of the
road was through brushwood, which had sprung up where the high forest had been
cleared for planting maize; but Belt soon found himself under a canopy of high trees
the trunks of which were entwined with creeping aroids and lianas, sending down their
: great rope-like stems to the ground. This forest is always wet, and the undergrowth
consists of small palms and magnificent tree ferns, with thin stems and delicate foliage,
and broad leaved heliconie, leathery melastome, and flesh-coloured begonias, with a
variety of other damp forest loving plants.

In 1872 Belt made a long journey to Segovia in order to engage labour, as the
Indian miners mostly came from that province. The road lay over a rough forest
country on the east side of the range dividing the great lake valley from Matagalpa,
and this part of his journey strongly contrasted with any former one, as he was now
at a long distance from the Atlantic, in a dry and arid region, due to the north-east
trade wind having deposited its moisture on the intervening stretch of high land.
Belt crossed several high ranges before reaching Ocotal, the capital of Segovia, situated
near the sources of the Rio Wanks; here grew pine trees and evergreen oaks at

F2

36 PHYSICAL FEATURES ETC.

probably the southern limit of the former in Central America. Descending a steep
slope beyond Ocotal, he came to a forest resembling that around Santo Domingo,
though the trees were not so large, but tree ferns, palms, lianas, broad-leaved heli-
coniz, and melastome were again abundant, and he was told that the Quezal, the
royal bird of the Aztecs, was occasionally met with. Belt, having successfully obtained
the required number of Indians, returned by nearly the same route to Santo Domingo,
and shortly after left for England.

Mr. C. W. Richmond, who resided in Eastern Nicaragua from February 1892 to
January 1893, when describing the climate, says [Proc. U.S. Nat. Mus. xvi. (1893) ]
that the east coast has a protracted rainy season of eight or nine months, with
occasional spells of fair weather; the rainfall is enormous, in some years reaching
296 inches at Greytown. He went up the Rio Frio into Costa Rican territory, from
the Lake of Nicaragua to the Guatusa Indian settlements, at the head of the canoe
navigation. A shark, doubtless the same species as that known to inhabit the lake
(Carcharias nicaraguensis), was seen as far up the river as he ascended. Later
Mr. Richmond spent some months on the Escondido River, chiefly about 50 miles
from the mouth. This river was formerly known as Bluefields, or Blewfields, and is
probably the most important on the coast of Central America, with the exception
of the San Juan. There is no troublesome bar, as is usually the case, and large ocean
steamers ascend to Rama, 65 miles from the mouth, where two rivers, the Rama and
the Sequia, join and form the Escondido. The banks for many miles, including both
branches above Rama, are Jined with banana plantations, the monotony of which is
broken by the numerous picturesque ceiba and ebo trees which have been left standing
in the clearings, while the dense tropical forest lies in the background. In the last
15 or 20 miles of its course, the river winds through dreary silico swamps nearly
devoid of bird life, and then empties itself into the Bluefields Lagoon, 15 miles
long and 7 miles broad. Mr. Richmond enumerates 281 species of birds which
he observed during this journey.

COSTA RICA.

This country was not visited by either Salvin or myself, and I am indebted to
Mr. Carriker, Mr. Ridgway, and other writers for the following information.

The little Republic of Costa Rica has an extreme length of 250 miles with a breadth
of about 150. ‘The greater part of the country is very mountainous, with narrow
coastal plains on both sides, finally extending to about 30 miles in width in the north-
eastern corner. ‘The drainage system is complicated, the extreme northern portion is
comparatively low, draining into Lake Nicaragua to the north and the San Juan
River on the Atlantic; while on the eastern side the streams and rivers have
their sources in the high mountains, and descend rapidly through narrow valleys or
gorges, separated by abrupt forest-clad ridges, which are frequently very narrow. The

COSTA RICA. 37

country is extensively wooded, the forests consisting of trees of all sizes, many of them
attaining enormous girth and height, especially on the eastern slope, where the rainfall
is most abundant; consequently, vegetation is there much denser and penetration very
difficult. On the Pacific side the forests have less undergrowth, the trees are larger
and taller, and progress through them is comparatively easy. ‘There is, however, an
exception in the Guanacaste region in the Nicoya peninsula, where great tracts of
erass lands or savannas, with scattered patches of woodland prevail. Trees with
berries and other kinds of fruit abound at all altitudes, furnishing food throughout
the year for the multitudes of tanagers, finches, parrots, toucans, and trogons, while
their blossoms give sustenance to innumerable humming-birds. The tree which,
throughout the tropics, mostly attracts other birds is one of the Leguminose,
bearing biennially great masses of fragrant tassel-like blossoms which persist for some
days before fading, while the season of flowering extends over a period of more
than two months; it is known as the ‘guava’ (this is not, however, the guayava of
commerce).

On the Caribbean slope the rainfall is fairly continuous during the whole year, reach-
ing on an average from 200 to 230 inches on the lower land. ‘The greatest rainfall is
from the middle of December to the middle of January, and again from the middle of
June to the middle of August, while from January 15th to March Loth it is fairly dry
and cool, and this season on the eastern slope is delightful. In the high regions the
rainfall is less, and there are alternately six months of wet and dry season. During
the first and last months of the rainy season the fall is slight and rarely of daily
occurrence, while in July, August, and October it rains every afternoon, and often
violently. During the dry season high winds prevail at all altitudes, but little if any
wind blows during the wet season. Influenced by the constant rain, the vegetation is
most luxuriant, and with it we find animal life consequently much more abundant.
So numerous are the species that probably not less than three hundred and ninety
land and fresh-water birds may be found on the lower portion of the Caribbean slope
up to 3000 feet. The high peaks rising above the regions of the Central plateau
constitute quite a distinct life zone. The main range of mountains extends from the
extreme north-west to the eastern central portion, gradually increasing in height and
ending in a chain of volcanoes reaching an altitude of from 8000 to 11,000 feet,
including Poas, Barba, Irazu, and Turrialba. Here there is a break formed by two
deep, broad valleys, the river Reventazon flowing in the one to the Atlantic and the
Rio Grande de arcoles in the other to the Pacific; their sources are within half a
mile of each other, the ‘divide’ or watershed being known as ‘ El Alto’ and having
an elevation of 5000 feet. The whole country to the south is an unbroken mass of
mountains containing but few inhabitants except the rapidly diminishing Indians.

Mr. Carriker (Ann. Carnegie Museum, vi. 1910) considers that the Avifauna of
Costa Rica is composed of three primal groups—the Boreal, the Sonoran, and the

38 PHYSICAL FEATURES ETC.

Tropical,—the first two coming down from the north, the last coming up from the
south, and all meeting in the Republic of Costa Rica. He says, and truly, that
the geographical position and meteorological peculiarities of the country make this
enormous bird fauna within its confines explicable, but at the same time they greatly
increase the difficulty of a satisfactory disposition of many of the species with respect
to life zones. The continent decreases from 3000 miles in breadth to scarcely more
than 60 at the narrowest part of Costa Rica, and within those 60 miles are crowded a
diversity of climatic conditions, altitudes, etc., scarcely paralleled in the world. ‘The
northern and southern forms of the Pacific and Atlantic lowlands meet here and
overlap, a single remnant of the Canadian bird-fauna persists on some of the isolated
peaks of the high mountains (Junco vulcani), while modified forms of this northern
species are found on the high lands of Mexico and Guatemala. A very large number
of North American migrants arrive during the winter months and distribute themselves
widely as to altitude throughout the country. Mr. Carriker also observes that amongst
certain species there is a seasonal migration from a higher to a lower altitude,
doubtless for the sake of food, and Salvin and I noticed similar migrations of
several species of birds in Guatemala; this was especially the case with humming-
birds, which were very abundant at Duefias during the flowering season, but in the dry
season, when the flowers failed, this district was entirely deserted and the birds
migrated to the Pacific Coast, where the atmosphere was damper and the blossoms on
which they fed were abundant.

The cultivation of bananas, however, is having a marked effect on the birds, which
are decreasing at an alarming rate, for in the few years since the establishment of the
Fruit Company at Port Limon the forest has been completely cleared—not only there,
but also southward towards the Talamanca district. Very few of the forest birds
frequent the banana plantations, and with the destruction of the forest they recede
or disappear altogether.

Mr. Ridgway (‘ Condor,’ vii. 1905) gives an interesting account of Poas, which is
the only active volcano in Costa Rica. After spending the night at San Pedro,
he resumed his journey by bright moonlight at 4 o'clock the next morning. The
‘lecheria’ or dairy farm at the upper edge of the cleared zone was reached soon after
daybreak, and the primitive forest which covers the last thousand feet of the mountain
was then entered. No pine trees were found here, as would have been the case at the
same altitude in Mexico or Guatemala, as south of Nicaragua they cease to exist.
The density of the forest was, however, such that it was impossible to leave the track
without cutting a way with ‘machetes,’ and as the undergrowth consisted mostly of
slender climbing bamboos, with exceedingly hard stems, which almost filled the
spaces between the trees, the difficulty of making much headway may be imagined,
‘Lhe variety of trees was very great, and all were laden with orchids, bromeliads, and
mistletoes, the latter often conspicuously and brilliantly flowered and the bromeliads

COSTA RICA. 39

of dazzling hues of orange, scarlet, and crimson. Here, too, was seen the Quezal, amid
surroundings no less magnificent than itself.

Leaving the horses in the open basin of an ancient volcano surrounded by forest,
Mr. Ridgway proceeded on foot to the summit of the cinder cone, but the view was
disappointing, as the crater was filled with dense clouds, except for one moment, when
a strong wind dispersed the mass of vapour and allowed a brief glimpse of the boiling
lake, 400 metres below. From the summit a descent was made to the lagoon, another
extinct crater filled with clear water almost icy in its coldness and surrounded by dense
forest. The time spent on Poas was too short to learn much of the birds, but except
at the summit they were everywhere found in great numbers.

At Bonilla, an estate on the Atlantic slope near Turrialba, the roughness of the
ground and the density of vegetation in the hot humid zone made it very difficult to
collect, but in the cleared areas, where the ‘potreros’ or pastures are found, the
variety and abundance of birds was remarkable. In one locality at least four hundred
species were obtained, and in the thick growth many escaped capture, while humming-
birds were so plentiful that fifteen species were shot round one flowering ‘ guava’ tree.
From here also was witnessed a flight of migrating hawks, passing northwards in
hundreds and thousands.

At Coliblanco, near Turrialba (6500 feet), the trees, including the brilliant scarlet-
flowering Lrythrina, were covered with creepers and epiphytes, while near the
water-courses at least three species of magnificent tree-ferns were common, as well
as Caladium-like plants with leaves large enough to form a shelter against rain.
As might be expected at this altitude, the birds were mostly different from those
of Bonilla.

Another interesting place was Pigres, at the mouth of the Rio Grande de Tarcoles,
in the Gulf of Nicoya. Between this and the mainland proper lies the ‘estero,’ a
broad creek of smooth water, bordered by dense mangrove swamps, somewhat narrow
on the Pigres side where the land mostly consists of bare sand, but in places covered
. with trailing [pomea, bearing broad leathery leaves and pink flowers; matted clusters
of thorny leguminous shrubs and thickets of low spreading mimosa-like trees inter-
_spersed with the poisonous manzanilla, formed the rest of the vegetation. Notwith-

standing the dry season and the almost total absence of flowers, birds were very
numerous in the vicinity of the village. Immediately beyond the fringe of mangrove
swamps, on the other side of the ‘estero,’ a high mixed forest extended for many
miles, the undergrowth consisting chiefly of small biscoyal palms bristling with ©
long slender thorns of needle-like sharpness. Further inland these palms gave place
to high cannas, and in the forest of tall trees, macaws, parrots, and parrakeets
were numerous and noisy.

An expedition from Coliblanco was made to the base of the cinder cone of Turrialba,
about 9100 feet. Here everything was different from what had been seen below.

40 PHYSICAL FEATURES ETC.

The several hundred acres composing the ‘potrero’ resembled an immense well
kept park, with long vistas through groves and clumps of magnificent trees on
undulating grassy slopes, cropped closely by the grazing cattle. On the right rose the
cone of the volcano, covered with dense ‘chaparral, or bush of evergreen oaks,
while to the left the long ridge-like mass of Irazu was plainly visible. :

Mr. Ridgway made an ascent of this volcano, about 11,500 feet. The forest, of
which but little remains, consists chiefly of oaks and differs widely from that of Poas
and Turrialba. Higher up and close to the ash-cone the trees become scarce and
scrubby, and finally only a growth of stunted Vaccinium-like shrubs exist.

It is remarkable that in Costa Rica at least 700 species and subspecies of
birds have been found. Dr. Outram Bangs, when alluding in the ‘ Auk’ (1907)
to the Costa Rican collections made by Mr. Underwood, remarks that the extensive
bird-fauna of this small country, scarcely larger than the State of Florida, is due
to the fact that the Central American forms extend to the Atlantic lowlands, while
those from Panama and the south go up the Pacific slopes, separated only by the
range of high mountains.

An account of the Costa Rican Odonata, their larval forms and their habits, is
given by Dr. Calvert in the ‘ Entomological News’ for July 1910. He and his wife
remained in the vicinity of Cartago for a year, making collections of Odonata as well
as of terrestrial molluscs, annelids, araneids, orthoptera, microdiptera, coleoptera, and
lepidoptera to a smaller extent, but they were hurried away by the severe and frequent
earthquakes which finally destroyed the town in May 1910.

PANAMA.

This State—or Republic, as it must now be called—comprises the neck of land
extending from Costa Rica to Colombia, an area equal in extent to about two-thirds
the size of England and Wales, and forming the most southern country dealt with in
the ‘ Biologia.’ Very little, however, of the Isthmus of Darien, the land south-east of
the Canal, has been visited by collectors. The main chain of the Cordillera decreases
greatly in height towards the City of Panama, and between that place and Colon, where
the railway and Canal traverse the country, the elevation falls to less than 300 feet.
Salvin crossed by rail on more than one occasion, and spent some time collecting near
the Station of Obispo, where he obtained a good many specimens ; but Enrique Arcé
and Mr. Champion were specially employed in Chiriqui, Arcé subsequently proceeding
to Veraguas where he remained for several years.

The rivers, taken as a whole, are unimportant, but the Chagres with its tributary the
Obispo attains formidable dimensions in the wet season, overflowing j
inundating a large area. ia eat tees Ares

The district immediately adjacent to the Canal has recently been described by Mr. A.

PANAMA. 4]

Busck (‘ Report on the Mosquito Fauna of Panama,’ 1908), who spent three months in
the neighbourhood of Tabernilla near Colon. The ground slopes towards the Chagres
River, and in the intervening country lies the bed of the old French sea-level canal
which, even in the dry season, is covered with a series of shallow lakes connected by
low marshes. Between these and the river are tall bamboos, sparsely interspersed with
large hardwood trees, the crowns of which are covered with parasitic plants, orchids,
and tillandsias, the last named affording a breeding place for several species of
mosquito.

When Mr. Champion visited Panama in 1881-1883 he investigated the Pacific slope
only, that on the Atlantic side being very inaccessible, and except at Colon and along
the railway, or near the coast, there were no villages or means of obtaining food or
shelter. He endeavoured to ascend the Volcano of Chiriqui, which attains an elevation
of 11,000 feet, but the only route through the forest lay by narrow tracks made by
tapirs, and on reaching the summit of a ridge, at 8000 feet, further progress was im-
possible, owing to the presence of an immense ravine, from which the upper part of the
volcano could alone be seen. The night was spent in a hut erected by orchid collectors,
but as no water could be obtained he was obliged to descend the following day.

On the western slopes of the volcano the savannas reach an altitude of about
6000 feet, while at 4000 feet cattle are pastured in large numbers for the Panama
market. Higher up, to the north and west, a dense belt of forest covers the mountain
side, but this does not extend to the summit. On the southern slope the forest had
been cleared in many places for the cultivation of coffee, and a fine palm was locally
abundant, but Coniferz were entirely wanting. At Chorcha (300 feet) the dense forests
descended to the coast. and interrupted the continuity of the large savannas bordering
the Pacific Ocean.

The Avifauna of Central America south of the Lake of Nicaragua, including Costa
Rica, Chiriqui, Veragua, and Panama, is exceedingly rich. These countries contain
more species than the whole of Europe, and nearly as many as the whole of America
north of Mexico; 432 species have been found in Veragua, including Chiriqui.

Areé’s collections, like those of other naturalists who have since visited the country,
were almost, if not entirely, made on the southern or Pacific slope.

Summarising his analysis of the birds of Veragua *, Salvin remarks (P.Z.S. 1870,
pp. 178-179) as follows :—“ The characteristic elements of the Central American fauna
consist not so much in the amount of generic peculiarity, which is very small, as in the
fact that a very considerable portion of South American forms are here represented,
not as specifically identical, but, in a large number of instances, as definably distinct
in degrees of varying value. The element of the Central American bird-fauna to
be traced to the northern continent, on the other hand, maintains a very different
relationship to the bird-fauna of that continent. With the exception of a few species

* Chiriqui was included by him under Veragua.

BIOL. CENTR.-AMER., Introd. Vol., January 1915. G

42 PHYSICAL FEATURES ETC.

isolated in the mountains of the higher portions of the Isthmus, and some others, we
find that northern forms found in Central America are specifically identical with
northern species, and that their presence is due in a great measure to migration during
the winter season. As regards numbers, we find a gradual diminution as we recede
from North America. These migrants are everywhere present, some few passing still
further south into the equatorial provinces of the southern continent. Costa Rica and
Veragua, with Panama, possess these characteristics of the Central American fauna in
the highest degree. It is here we find the greatest number of South American genera
represented; but the species are to a considerable extent not the same as the
_ continental species. If we endeavour to account for the facts as we find them, by
changes in past times in the physical features of the Isthmus, we seem to require :—
1st. A union between Costa Rica, Veragua, and Panama with the southern continent,
when those united lands possessed in common a much larger number of species
specifically the same as at present, during which time the oceans may have been
united north of Costa Rica. 2nd. The long duration of Costa Rica and Veragua
as a ‘continental’ island, when the union of the two oceans has been of greater
extent. This period must be long enough to have established specific differences
much as we now find them. 38rd. The emergence of the whole Isthmus in its present
form. These requirements seem to fall in fairly with what has been demanded in
other branches of natural science. Dr. Duncan requires a union in Miocene times
between the oceans to account for the specific identity of certain corals. The union
here demanded will suit my first and second requirements, I only regulate the amount ;
and as for the period when it took place, fixing it to Miocene times would seem to
answer to the requirements of the birds. That all the peculiar features of so varied
a fauna can be accounted for by this theory I do not pretend to say. The changes in
the physical features of the Isthmus, indicated by the numerous minor modifications
of existing species, belong to the most recent events in geological history. To
account for the greater differences observable we must go deeper into the abyss of
geological time, where light is at present barely perceptible.”

In his first paper on this subject (P.Z.S. 1867, pp. 129-161), based upon less
extensive material, Salvin stated that there was a closer affinity between the birds of
Veragua and those of Costa Rica than between those of Veragua and of the Isthmus
of Panama, but this proved not to be the case when Arcé’s later collections were
examined. He then remarked that it was evident that Costa Rica and Panama had
for a long period occupied the position of one or more islands between the two
continents at a time when the two oceans were united by two or more channels; and
that an obvious division separating Costa Rica, Veragua, and Panama from the
southern continent was a line drawn from the Atlantic Bay of San Blas to the mouth
of the Bayano on the Pacific.

PEARL ISLANDS.. 43

PEARL ISLANDS.

In the Bay of Panama there lies a small group of islands known as Las Perlas,
or the Pearl Islands, consisting of Pedro Gonzales, San José, and San Miguel; the
last mentioned, also known as the Isla del Rey, is by far the largest, only twenty
miles from the nearest mainland and sixty from Panama. This island is about fifteen
miles long and irregularly oblong in shape, covered with low hills, which in turn are
clothed with luxuriant tropical forest. The climate is hot and unhealthy, and the
population consists almost entirely of negroes, who manage the affairs of the island
and are very independent of the Panama Government. The pearl-diving industry
having been almost abandoned, the people now grow vegetables, coco-nuts, and fruit
for the Panama market.

Mr. Champion, who visited San Miguel on our behalf in April 1883, was only there
in the dry season, and the “luxuriant forest” of other writers he describes as
“scrubby wood.” ‘The interior was somewhat inaccessible, the few paths or tracks
leading only to the patches of cultivated ground. The coast, like that of the
adjacent mainland, is covered with mangrove-swamps, which can only be traversed
at low water. | Pa

The Islands are in such close proximity to each other that probably the birds would
be the same on each. On San Miguel 46 species were found, of which only four
were considered by Mr. Bangs (‘ Auk,’ xviii. pp. 24-33, 1901) to be well-marked
island-forms, the remainder were, as might be expected from the semicircular form
of the Coast of Panama, similar to those of the adjacent mainland. Some birds are
undoubtedly carried across to the Islands by storms, but others perform the journey
voluntarily, among them a small green humming-bird (Chloristilbon assimilis), which
has been seen in perfectly calm weather flying straight for the Archipelago.

Certain well known butterflies from the mainland also occur, including a Morpho
( peleides 2), etc.

[ 44 ]

SOURCES FROM WHENCE OUR MATERIAL WAS OBTAINED.

Tue following particulars as to the sources from whence our material was obtained
supplement the account given in the Introduction, antea, pp. 11,12. In Volumes I.
of the ‘Aves’ and of the ‘Lepidoptera Rhopalocera’ the names of the various
collectors who were specially employed to obtain material for the present work are
recorded, viz. Messrs. F. B. Armstrong, E. Arcé, G. C. Champion (1879-1883),
W. Lloyd, W. B. Richardson (1897-1898), H. Rogers (1877), and M. Trujillo (1888),
and Mr. and Mrs. H. H. Smith (1888). From many others, also, we acquired or
received vast numbers of specimens, all of which were duly acknowledged by our
Contributors.

As regards the Mammalia, Aves, Reptilia and Batrachia, Pisces, Mol) usca,
Lepidoptera Rhopalocera, and Neuroptera (Odonata), the sources from whence our
collections were derived are mentioned in detail in the ‘ Introductions’ to the volumes
on these subjects and need not be repeated. The Coleoptera, or beetles, however,
numbering 18,029 species in all, and requiring eighteen volumes for their enume-
ration, by twelve contributors, demand special notice. Notwithstanding their vast
number, and the great abundance of individuals, this is the only group of insects,
the Lepidoptera excepted, for which sufficient contributors have been forthcoming to
enable us to complete their investigation. The Coleoptera, therefore, occupy in this
work an undue amount of space amongst the thirty-eight completed volumes of
Insecta, as compared with the Hymenoptera, Neuroptera, &c., certain important
families of which still remain unworked, this being due to the fact that the beetles
have proved more attractive both to the collector and describer.

The Coleoptera examined have been mainly supplied by the following col-
lectors from each of the various countries :—Mexico—A. Sallé, C. T. Hége (who
collected in nearly all the different States, Tamaulipas and Yucatan excepted), H. H.
Smith (who specially visited Guerrero, Morelos, Vera Cruz, Tabasco, &c., for us),
G. F. Gaumer (Yucatan), A. Forrer (N.W. Mexico, including the Tres Marias Is.),
J. Flohr, F. D. Godman (Central and Southern Mexico), M. Trujillo (S. Mexico),
Becker (Durango), Buchan-Hepburn (Chihuahua and Durango), and H. F. Wickham ;
British Honpuras—F. Blancaneaux ; GuaremaLa—A, Sallé, O. Salvin, J. Rodriguez,
G. C. Champion, L. Conradt, and F. Sarg; Nicaragua—lT’. Belt, E. M. Janson, and
W. B. Richardson ; Costa Rica—H. Rogers, P. Biolley, Van Patten, C. F. Underwood,

SOURCES OF MATERIAL. 45

and H. Lankester ; Panama (including Chiriqui, the Pearl Is., and Taboga)—G. C.
Champion, EK. Trotsch, H. Ribbe, and A. Boucard. The Hymenoptera examined
and reported upon by P. Cameron mainly consisted of the collections made by
G. C. Champion in Guatemala and Panama, and by G. F. Gaumer in Yucatan, most
of the vast number of specimens obtained in Mexico by H. H. Smith having been
received too late to be included; it must be remembered, too, that the bees and social
wasps have not been dealt with. ‘The Lepidoptera, apart from the specimens captured
by Salvin or myself, were obtained from the collectors who supplied us with the other
insects; very extensive additions to both the Rhopalocera and Heterocera have,
however, been made since 1900, especially by W. Schaus, who visited Mexico and
Costa Rica in search of them. The Diptera examined were comparatively few in
number, though H. H. Smith did his best in Mexico to make up the deficiency. The
Rhynchota were very numerous, both in Heteroptera and Homoptera, as regards
Guatemala and Panama, but till H. H. Smith went to Mexico we had received
very little from that country, the habitat of most of the previously described
species from our region. The Orthoptera, again, were few in number, none of our
collectors having paid very much attention to them. The Arachnids were mainly
supplied by H. H. Smith (Mexico), F. Sarg (Guatemala), and G. C. Champion
(Panama); the Acaridea, however, were mainly procured by, and belonged to,
Dr. Otto Stoll. In the Botany, the collections made by Mr. and Mrs. Salvin were
used by Mr. Hemsley, but the study of the plants was almost entirely made from
material contained in the Herbarium at Kew, including that which we had previously
sent from Guatemala. Mr. Maudslay’s work on the Archeology was based on his own
observations on the various ruins visited during his sojourn in the country; Mr. J. 8.
‘Goodman, in his Appendix to this subject, gives an account of the system he used to
decipher the Archaic Maya Inscriptions, which relate to a series of calendars. |

Mr. Champion’s Itinerary is given on pp. 46-54, and the places he visited in
Guatemala are easily traceable on Map 8; we are unable, however, to find space
on our other maps to include all the Mexican localities quoted in the Zoology and
Botany, some of which, indeed, cannot now be traced *.

* It must be remembered that nearly all the names commencing with ‘San,’ ‘ Santo,’ and ‘ Santa’ have
been used over and over again in the different States. The spelling of some of the others has been emended

on our Maps to agree with official Mexican diréctories.

[ 46 ]

ITINERARY OF
MR. G. C. CHAMPION’S TRAVELS IN CENTRAL AMERICA,

1879-1883.

THE only detailed itinerary supplied by any of our collectors is that of Mr. G. C.
Champion, who, it must be remembered, devoted almost the whole of his time
to entomology. A brief account of the numerous Guatemala and Panama localities
he visited will be of interest to entomologists. His expedition to Central America
extended from March 16th, 1879-May 28rd, 1883. The itinerary of his travels *, which
is here reproduced with additional details, appeared in the American ‘ Entomological
News’ for February 1907, pp. 33-44. The Guatemala routes are shown on Map 8.
During his stay in that country, March 16th, 1879-April 7th, 1881, he investigated
both the Pacific and the Atlantic slopes, the central plateau, &c. From August 10th,
1879-July 26th, 1880, he made his headquarters at San Gerdnimo, six miles from
Salama, and made many distant expeditions to various parts of Alta and Baja Vera Paz
from that place. The central plateau was traversed from Salama (vid Rabinal,
Cubulco, Joyabaj, and Quiché) to Quezaltenango, and also from the latter place to the
Capital. The Pacific slope was explored from La Gavia to the Mexican frontier of
Chiapas, at elevations between 1000 and 4000 feet. The ports of San José and
Champerico, the lagoons at Paso Antonio, and the lakes of Amatitlan and Atitlan
were visited. The Volcan de Agua was ascended on January 13th, 1881. In Vera
Paz, the Rio Dolores, near Cubilguitz, on the Coban—Peten road, was reached; and
the Polochic Valley was twice visited, once by boat from Panzos as far as the Lake of
Yzabal. An expedition was also made across the Chuacus range of mountains from
San Gerénimo to Tocoy in the Motagua valley. Mr. Champion is of opinion that,
from what he saw of the country during his travels, Guatemala might well be divided
for zoological or botanical purposes into three (or four) districts :—(1) “The Atlantic
slope,” which is mainly of limestone formation in Alta Vera Paz, and has a very long
rainy season—in Coban sometimes lasting into March,—-and an abundance of humid

* All made on horse- or male-back or on foot; the first Guatemalan railway—from San José to Escuintla—
was opened just before he left.

ITINERARY. 47

forest; (2) ‘The Central Plateau,” including the ‘ Altos,’ which is an arid upland
region becoming higher towards the Mexican frontier and bordered southward by.
volcanoes which protrude from the main cordillera—pines and oaks here clothe the
mountain sides, on which cereals, maize, and, at high elevations, potatoes are culti-
vated ; (3) “The Pacific slope,” which has forest in the ‘tierra caliente’ and on the
mountain sides, now largely replaced by second growth (rastrojo) or cultivated with
sugar-cane, cacao, or (at elevations up to 4500 feet) coffee. The plain of Salama, in Baja
Vera Paz, with San Gerdnimo at its eastern end—draining to the Atlantic,—is very
hot and arid, abounding in Cacti, Yuccas, Agaves, &c., like the Motagua valley, from
which it is separated by the Chuacus range of mountains. The mountains to the north
of this plain, as shown by the abrupt change in the nature of the vegetation between
Salama and Purula or Tactic, form the dividing line between the humid Atlantic slope
and the plateau. These lower central valleys must either be included under district 2
or treated as a separate faunal subregion. A comparison of the Lepidoptera Rhopalocera
alone illustrates the great difference in the fauna of the two slopes *—many Erycinids,
Ithomiids, Heliconius, Papilio, Leptalis, Thecla, &c., are peculiar to the Atlantic ;
while Drucina, Euterpe, a few special Euptychia and Heliconius, a Morpho, &c., are
found on the Pacific. The dry central plateau doubtless forms an impassable barrier
for many species, and it has altogether a very restricted butterfly fauna. In the
mountains in the neighbourhood of the plain of Salama a few peculiar Rhopalocera
occur, such as Anewa nobilis and eacellens, &c. The ‘tierra fria’ or * tierra helada’
(10,000 feet and upwards) produced no alpine or subalpine forms, merely stragglers
from below. |

In Panama, April 17th, 1881—May 21st, 1883, Mr. Champion spent nearly all his
time in Chiriqui, on the Pacific slope, between the Rio Chiriqui Viejo (near the Costa
Rica frontier) and Tolé, making his headquarters at the various coffee-plantations on
the mountain-slopes, at an elevation of 3000-4000 feet, or else at David or Bugaba.
The Volcan de Chiriqui was ascended on June 7th, 1882, to 8000 feet, and the
Cordillera above Tolé explored. ‘The old route across the mountains from David and
Caldera to the Chiriqui lagoon and the Bocas del Toro Islands, on the Atlantic side
(used during the early days of the gold-mining in California, and before the Panama
Railroad was finished), was found to be almost impracticable, and the northern slope
was therefore not visited. The principal forests in Chiriqui are situated on the mountain
slopes, in the low country to the east of David, and in the‘ tierra caliente’ to the west of
Bugaba and Divala; the forests alternate with extensive savannas along the lower part
of the Pacific slope and in the country immediately adjacent to the western precipices
of the Volcan de Chiriqui. The whole of the towns and villages are situated in the
‘tierra caliente,’ and the Indians living in out of the way places in the Cordillera

* Unfortunately this could not be very well shown in the Table of Distribution of the genera given in the -
Introduction to the Rhopalocera.

4§ ITINERARY.

or on the northern slope are very seldom seen. On the southern slope of the Volcano,
between 2000 and 4000 feet, a great deal of the forest had already been cleared (in
1881) to plant coffee. San Miguel (Isla del Rey) in the Pearl Islands and Tobago were
visited by Mr. Champion, in April and May 1883, from Panama, and a certain number
of insects, &c., were collected by him in these places. The absence of Conifers in the
mountains, the paucity of Cacti, and the much less arid nature of the country afford
a striking contrast to Guatemala, the fauna of Panama being very similar to that
of Tropical South America. There is no arid central plateau in Panama, and the
Cordillera, the loftiest part of which is in Chiriqui, decreases towards the isthmus,
where it is only a few hundred feet high, so that the fauna of the two slopes is not
likely to differ greatly. ‘he Atlantic slope, however, bas not yet been investigated. A
Tenebrionid-beetle of the seashore, Phaleria dytiscoides, is recorded by Mr. Champion
as common to the coasts of British Honduras on the Atlantic and of Guatemala
and Nicaragua on the Pacific [ef. Coleopt. vol. iv. pt. 1, pp. 218, 219 (1886)],
indicating a former connection of the two oceans at the isthmus of Panama. His
itinerary was as follows * :—

1879. March 16-18. San José de Guatemala, the Pacific
port of arrival for travellers from Panama or San
Francisco. Sea-coast, mangrove-swamps, lagoons, &c.

March 19, 20. Travelling up to capital vid Escuintla,
by diligence, over execrable roads, all inches deep in
dust at this (dry) season.

March 21-April 2. Guatemala city (about 4500 feet).
Open plains, intersected by deep barrancas (ravines).
Scrubby oak and pine woods in places on_hill-sides.
The volcanoes Pacaya, Agua, and Fuego visible to

1879. May 13-June 22. Zapote (about 2000 feet). On the
forest-clad southern slope of the Volean de Fuego.
Broad, deep, dried-up watercourses, full of great
boulders, ran downward through the forest here,
making travelling difficult. The locality good for
insects. Coffee cultivated. Some very fine forest
passed through at San Cayetano, between Zapote
aud Capetillo, along the descending coast-read.
June 24, 25. Antigua.
June 26-July 21. Dueiias (about 4500 feet). Near

the southward. Many insects found on the banks
of the streams in the barrancas.

April 3-5. Ciudad Vieja. The first capital of Guate-
mala, in the valley between the volcanoes Agua and
Fuego. Coffee-plantations and cultivated ground,
unsuitable for collecting-purposes.

April 6-8. Guatemala city.

April 9. Aceituno. Coffee-plantation near the capital.

April 10-16. Guatemala city.

April 17-May 12. Capetillo. Valley between the
voleanoes Agua and Fuego. Coffee and sugar-cane
plantations, with the forest-clad slopes of the Fuego
adjacent. This estate is the property of the well-
known Guatemalan naturalist, Juan J. Rodriguez,
who has, from time to time, fur upwards of thirty
years, supplied the editors of this work with material
from his district.

Capetillo. Coffee and Opuntia (for rearing the
cochineal-insect) plantations adjacent to the Lake
of Duefias. Various excursions made from here to
Calderas, on the upper eastern slope of the Volcan de
Fuego, up to about 7500 feet. Pines on the higher
slopes. Earthquake-shocks frequent—the house be-
longing to the owner of the estate (who had to leave
it and live in Antigua) in ruins. The Acatenango
peak of the Volcan, as well as the smoking Fuego
itself, conspicuous from Dueiias.

July 22-Aug. 7. Guatemala city.

Aug. 8. Carrizal. Arid district with scrubby woods.

Aug. 9. Llano Grande (about 2600 feet). Scrubby
woods, cultivated ground, and pasture. Mule-trains
and Indians bearing heavy loads constantly met with

here, the route from the capital to Salama, Coban,
&c., passing through this place.

* Elevations approximate only, taken from an uncorrected aneroid barometer.

Probably too low in many cases.

ITINERARY.

1879, Aug. 10-Sept. 9. San Gerénimo, Baja Vera Paz (about

2950 feet). East end of plain of Salama, and six
miles distant from the town of that name. Hot,
dry region, with many cacti. Chuacus range of
mountains adjacent, bordering the plain southward ;
lower slopes clothed with pines, with forest of
deciduous trees above. Sugar-cane and coffee planta-
tions near village, belonging to English owners, the
estate having a local reputation for the quality of
the aguardiente (rum) and sugar produced by them.
Drainage to Atlantic. Headquarters for about one
year. Many long excursions made from here to
distant places in both Alta and Baja Vera Paz, on
the Atlantic slope. Some fine butterflies (Anca spp.)
peculiar to the Chuacus range. Various Longicorn
(Ochresthes), Buprestid (Acmeodera), and other genera
of Coleoptera characteristic of the drier portions of
Mexico occur on the plain of Salama, A large
Buprestid (Chalcophora virginiensis) in the pines on

1879.

49

down to near Rihaco and upward towards Matanza.
Road from Sabo descending very abruptly through
forest in which a transparent-winged Pierine-butterfly
(Dismorphia fortunata) was abundant.

Oct. 16. Purula.

Oct. 17. Cachil. Open arid mountain-slopes north of
the plain of Salama, with agaves, palms (Zhrinax),
&c. <A peculiar Kuptychia (rubricata) taken here. *

Oct. 18—Nov. 3. San Gerdnimo.

Noy. 4-6. Tocoy (about 2000 feet), Arid district on

_ the Zacapa road.

Nov. 7. El Jicaro, near the Village of Guacamaya, on
eastern slope of Chuacus range. Scrubby woods,
pines above.

Nov. 8-12. San Gerdnimo.

Nov. 18,14. Purula.

Nov. 15. San Miguel Tucuru (about 2000 feet).
Polochic valley. Mostly cultivated ground, cotton,
maize, &c.

the Chuacus slopes. An Ithomiid butterfly (Dircenna Nov. 16. La Tinta. Polochie valley. Tropical vege-
klugi) seen swarming in the shady garden of the tation. Indigo formerly cultivated here, hence the
hacienda. name.

‘Sept. 10. Santa Barbara (about 4450 feet). Moun- Nov. 17-23. Senahu (2800 feet). Limestone moun-
tainous region east of San Gerénimo. Scattered tains north of the Polochic valley. Humid forests,

woods. Pines below. Cultivation of maize, &c.

Sept. 11,12. Santa Cruz (5500 feet). Mountainous
region of Chilasco, the watershed between the
Motagua and the Polochic, east of Santa Barbara.
Scattered woods of Liquidambar, &c. Forest appa-
rently all cleared in vicinity of village. Nights very
cold here, ~

Sept. 18-Oct. 1. San Gerénimo.

Oct. 2-5. Purula (about 4000 feet). Open ground
with a humid virgin forest adjacent on mountains.
A new track through the forest towards Cerro Verde,
an excellent entomological locality, the road to Sabo
also productive. These localities again visited in
April or May, 1880. The ‘quezal’ (Pharomacrus
mocinno) not rare in the dense forest on the Cerro
Verde road, and a ‘Howler’ (Mycetes villosus)
frequently heard between Purula aud Sabo.

Oct. 6, 7. Sabo (2900 feet), Clearing made in dense
humid forest to plant coffee, on a steep mountain-
slope. <A very productive locality. Pulea irritans,
however, swarmed to such an extent in the disused
hut used for sleeping-quarters that it was impossible
to remain very long in the place.

Oct. 8-15. Panimd * (1800 feet). Hot, narrow valley
of the Rio Sinanja, a tributary of the Polochic.

1880.

cleared in many places for coffee-plantations. Long
rainy season, but water scarce, rapidly disappearing
underground. <A district rich in land-shells.
visited in June, 1880.

Nov. 24-30. San Juan (1800 feet). A small coffee-
estate, no village, on the mountain-slopes north of
the Polochic. Forest mostly cleared. Mountains
of the Republic of Honduras visible to the S.E.

Dec. 1, 2. La Tinta.

Dec. 3-7. Tamahu, a few miles higher up’ the valley
than Tucuru (about 2250 feet). Mostly cultivated
ground.

Dec. 8. Santa Rosa (about 4000 feet).

Dec. 9-28. San Gerénimo.

Dec. 29. Tactic (4300 feet). Scattered Liguidambar
and other deciduous trees. Forest all cleared to near
the inaccessible mountain-tops.. Large Indian popu-
lation in district, hence the continuous clearing of
the trees to plant inaize (‘ milpas’).

Dee. 30, 31.) Coban, Alta Vera Paz (about 3800 feet*).

Jan. 1, 2. } Humid region, rainy season sometimes
extending into February or March.
cleared to plant coftee, maize, &e. Numerous German
traders and planters settled here. The residence for
many years of a keen zoologist, F. Sarg. Large Indian

Again

Forest nearly all

Valley, mostly cultivated with maize, &c., followed population.
Jan. 3. Tactic.
* Misprinted ‘ Pancina’ on many of the labels attached to insects in the Jan. 4-6. San Gerénimo.

collection made by Mr. Champion, and therefore wrongly quoted in some
of the volumes of this work.

* Maudslay makes it 4280 feet.
BIOL. CENTR.-AMER., Introd. Vol., January 1915. Hl

50 ITINERARY.
1880. Jan. 7. Buenaventura.
Jan, &-12. Guatemala city.
Jan. 13. Lake of Amatitlan (about 3450 feet). Arid

district, with many ‘nopales’ (plantations of Opuntia
for rearing the cochineal insect, all enclosed within
dusty adobe walls). The Volcan de Pacaya not very
far distant.

Jan. 14-18. Guatemala city.

Jan. 19. Buenaventura.

Jan, 20-25. San Gerdnimo.

Jan. 26. Santa Rosa.

Jan. 27—Feb. 1. San Joaquin, Alta Vera Paz (about
3200 feet). Pine-clad, arid mountain-slopes. Rio
Chisoy (or Chixoy) below. About the northern limit
of the arid region of the central plateau.

Feb. 2. San Cristobal (4250 feet). Cultivated ground
adjacent to the Lake of San Cristobal.
Feb. 3-5. Balheu (Valeu) (8850 feet),

mountain-s!opes.

Feb. 6-10. Coban.

Feb. 11. Chiacam (2400 feet).
scrubby woods.

Feb. 12. San Agustin Lanquin (1000 feet).
stone formation. Second-growth woods; forest all
cleared. Rio Cahabon, a large tributary of the
Polochic, adjacent. An unproductive locality.

Feb. 13-23, Cahabon (Cajabon) (about 800 feet).
Second-growth woods, forest all cleared near village.
Large Indian population. No other people here,
except the priest (who kindly accommodated stray
travellers) and two or three Guatemalan officials.
The very large church here is placed on a hill, higher
than the smaller ones on which the Indians have
built their huts. Plantations of cacao, maize, &c.,
around the village. ‘Toucans seen in these places.

Feb. 24-28. Lanquin. The large limestone cave
visited *, in company with an Austrian plant-collector
casually met in the village. Only insects seen in the
cave, an apterous Orthopteron (Arachnomimus
cavicola). The Rio Cahabon, a broad stream, issues
from its mouth, making its first appearance above
ground at this place.

Feb. 29-March 6. Chiacam.

March 7-9. Coban.

March 10-20. Cubilguitz (1050 feet), near Choctum.
Limestone region, with humid forest on the hills, the
roads between the hills extremely bad. Broad valleys
with scattered trees. The Rio Dolores reached, but
mot crossed. On main road from Coban to Peten,
the track to Salinas (a place where salt is obtained
and sent on Indians’ backs to Coban) turning off to

* Also entered by Salvin, on Mareh 8th, 1860 [cf ‘ Ibis,’ iii, pp. 140, 141
(i861)},

Pine-clad

Coffee-plantations and

Lime-

1880.

the westward near Cubilguitz. A good locality, but
food absolutely unobtainable from the scattered
Indian residents.

March 21. Satchicht (2000 feet). In fording the
river here my mule was so badly cut about the legs
by the jagged knife-edged submerged limestone
ledges that it was unfit for work for three months
afterwards.

March 22-24. Coban.

March 25. Tactic.

March 26-April 13. San Gerénimo.

April 14-23, Purula.

April 24-28. Panima.

April 20-May 2. Sabo.

May 3. San Miguel Tucuru.

May 4-12. Chacoj, near Chamiquin, sometimes called
La Hamaca (from the old suspended rope-bridge over
the Rio Polochic) (about 500 feet). Tropical forest,
with many palms (mainly Attalea cohune), which
decrease in size as the Polochie is ascended. A
new iron bridge in course of construction (in 1880).
A very good entomological locality.

May 13-18. Teleman, on the Rio Polochic, the upper
limit of navigation for small boats from the Lake
of Yzabal. Tropical forests, with many: palms, the
leaves of the lofty Attalea cohune arching across the
road. Culicide swarming. Unhealthy district. The
‘ Howler’ frequently seen in the trees near the river.
Some peculiar Syntomid- moths found amongst the
prickly herbage.

May 19-22. Panzos, on the Rio Polochie.
forests, &c., as at Teleman. Willows on river-bank.
Unhealthy district. My first attack of fever here.

May 23. Danta, on northern shore of the Lake of
Yzabal, about fourteen hours’ journey by small boat
from Panzos. Many alligators and ‘ Howlers’ seen
on my way down, manatees also noticed in the lake.
Culicide swarming.

May 24-25. Travelling up the Rio Polochic, two days,
against stream.

May 25-June 2. Panzos. Culicidee (especially a sooty-
black species) so bad here, even by day, that it was
almost impossible to do more than a few hours’
collecting at a time. Local name for them,
‘zancudos.’

June 3-14. Senahu, travelling up from Panzos by way
of Trece Aguas. [The American entomologists,
Messrs. H. 8. Barber and E. A. Schwarz (of the U.S.
National Museum at Washington) have visited this
district during recent years, and they succeeded in
obtaining various minute Coleoptera of the same
species captured by myself in 1880.]

June 15-21. San Juan.

Tropical

1880.

ITINERARY.

June 22, 23. Chaco}.

June 24. San Miguel Tucuru.

June 25-27. Purula.

June 28-July 26. San Gerénimo. My last (8th) visit
to this place.

July 27. Rabinal (2850 feet), en rowte for the ‘ Altos.’
Dry region, with scrubby woods, cacti, agaves, yuccas,
&e., as on the plain of Salama.

July 28. Cubulco (2900 feet). Similar country.

July 29. Joyabaj (4300 feet), an Indian village, reached
by a long precipitous ascent from Cubuleo. Open
mountainous region, intercepted by deep barrancas.
Pines and oaks on slopes. Central plateau becoming
higher westward. An interesting butterfly (Chryso-
phanus pyrrhias) seen in numbers on the way up from
Cubulco.

July 30. Santo Tomas Chiché (6100 feet). Los Altos
region.

July 31-Aug. 5. Chimente, Quiché Mountains (7600
feet). Indian village. Oaks, pine, alder, &c., on
slopes. Potatoes and maize cultivated. Cyanide
bottles (with the results of a day’s collecting) stolen
trom saddle-bags here.

Aug. 6. Totonicapam (7900 feet). Pine-clad slopes,
but too far from the town for collecting purposes.
Aug. 7. Chevuc (9900 feet). Pine woods. Be-
nighted at this place, having missed the road along

the Cordillera to Los Encuentros in the dark.

Aug. 8. Los Encuentros (8400 feet). A resting-
place for the night, on the road to Solola or the
capital. Arid open ground. ([Conradt collected
insects at Tecpan, a place to the eastward. ]

Aug. 9, 10. Desconsuelo (Solchicha) (about 10,500
feet). Pine-forests. Very bleak situation above
Totonicapam. Carriage-road from Quezaltenango to
Guatemala city passes this place, following the
summit of the highest portion of the Cordillera.
Potatoes only cultivated. A very gvod locality *,
most of the Coleoptera, and some of the Lepidoptera
Heterocera, collected proving to be new, but no
peculiar butterflies were met with.

Aug. 11-15. Pachoe (or Patchoc) (9200 feet). Indian
village. Pine-woods. Only accommodation obtainable
a small schoolroom used during the day, the bare
earth covered with pine-branches serving as a bed.
Slopes of the Cordillera accessible from this place.

Aug. 16. 'Totouicapam.

Aug. 17-19. Quezaltenango (7600 feet). Cultivated
ground mostly. Large Indian population. Un-
productive locality.

* The insects from Desconsuelo and Pachoe were collectively labelled
‘Totonicapam ’ (a town at the foot of the Cordillera) in the collections
made by Mr. Champion.

ima

51

1880. Aug. 20-Sept. 9. Finca of Las Nubes on the southern

(Pacific) slope of the Cerro (or Volcan) Zunil, above
Mazatenango (4050 feet). Extensive coffee-planta-
tions, with dense forest above. The most productive
locality visited on the Pacitie slope. Several new
butterflies (species of Drucina or Euptychia) met with,
An interesting bird (Oreophasis derbianus), a monkey
(Ateles ater ?), &c., seen in the forest. An enormous
Passalid-beetle (Proculus goryi) found commonly
beneath the large tree trunks left on the ground to
decay in the ‘ cafetales ’ (coffee-plantations).

Sept. 10-23. San Isidro (1600 feet). ‘Tierra caliente,’
Pacific slope, below Mazatenango. Second-growth
woods, cleared in places for coftee and cacao planta-
tions. A Cualigo plentiful in the tangled under-
growth, but difficult to secure.

Sept. 24. Retalhuleu (950 feet). Similar country.
Now connected by rail with the port of Champerico.
Sept. 25-Oct. 7. Las Mercedes (3200 feet). Pacitic
slope. Immense coffee-plantations in this Costa
Cuca district. Nearly all the original forest cleared,

Oct. 8-17. El Reposo (800 feet). Low country near
Pacific. Mostly second-growth woods, but some
forest, with lofty palms, in vicinity. Macaws
(guacamayo’s) often seen.

Oct. 18, 19. Paraiso(300 feet), Near Pacific. Scrubby
woods, bamboos, &c.

Oct. 20-22. Champerico. Sea-coast. The remarkable
fish (Anableps dovit) seen in the lagoons, [Also met
with by Salvin, at Chiapam, near here. |

Oct. 23. El Reposo.

Oct. 24-26. Las Mercedes.

Oct. 27, 28. Coatepeque (1250 feet). Mostly culti-
vated ground. [Village almost completely destroyed
by earthquakes during recent years. |

Oct. 29. Rio Naranjo (450 feet). Second-growth
woods. Mexican frontier adjacent. A new iron
bridge in course of construction.

Oct. 30. Near Naranjo. Benighted by losing road.

Oct. 31-Noy. 38. Finca La Union (2250 feet). El
Tumbador district, department of San Marcos.
Coffee-plantations and second-growth woods. No
villages hereabouts, the town of San Marcos on
higher ground in the Cordillera.

Noy. 4-7. Finca La Carolina (2600 feet). Tumbador
district, overlooking lower portion of Soconusco,
Chiapas, to Mexican coast, the Volcanoes Tacana
and Tajumuleo visible to the north-west. Unpro-
ductive locality.

Nov. 8. La Union.

Noy. 9. Rio Naranjo.

Nov. 10. Coatepeque.

Noy. 11-14. Las Mercedes.

H 2

1880,

52 ITINERARY.

Nov. 15. San Martin, near Ostuncalco (7400 feet).
Scattered trees, ground mostly cultivated.

Nov. 16-18. Quezaltenango. Cerro Quemado visited.

Nov. 19-Dee. 14. Las Nubes (Cerro Zunil). [The
village of Santa Maria, on way down from Quezal-
tenango, almost destroyed during recent years by an
eruption of the volcano of that name. |

Dec. 15. San Isidro.

Dec. 16-26. San Agustin (2250 feet). Southern
(Pacific) slope of the Volcan de Atitlan. Second-

growth woods, coffee-plaritations, &c., all very dry
and dusty at this season. Many Lepidoptera
Heterocera taken ‘at light’ in the verandah of the
house.

Dec. 27-29. San Lucas Toliman (4900 feet). Indian
village on the borders of the Lake of Atitlan, which
is very deep and has no visible outlet to the Pacific.
Oak-woods, &c., arid region. Pampojilaj, on the
coast road, visited, and the lake crossed (at night) to
northern side. Numerous thickly populated Indian
villages round the lake, the Indians not very friendly.

Dee. 30. Panajachel (4900 feet). Stayed at a flour-
mill, close to the lake. The town of Solola not very
far distant.

Dec. 31. San Lucas Toliman. Returned from Pana-

jachel by a tortuous detour along the high ground
above the lake.

. Jan. 1. Godines (6900 feet). Arid district above the

precipitous cliffs bordering the Lake of Atitlan on
the N.E. side. Magnificent view at sunset of the
lake and the adjacent volcanoes and mountains to the
southward during the cloudless skies of the dry
season.

Jan. 2. Chimaltenango (5650 feet). Reached by way
of Patzun and Patzitzia. Upland district cultivated
with cereals.

Jan. 3-11. Guatemala city.

Jan. 12. Antigua.

Jan. 18. Volcan de Agua (about 13,000 feet) ascended
at night (moonlight) from the Indian village of Santa
Maria (6500 feet). Belt of deciduous trees above
the cultivated ground to about 9500 feet, scattered
pines above, even in crater. Path up deep and
extremely narrow, between dense tussocks of high
grass. Very few insects met with on summit, the
butterflies seen merely common stragglers from
below. Indians ascend to fetch a little ice obtained
from holes made in the ground. So windy that it
was impossible to remain long on summit, clouds
forming rapidly after about 10 a.m.

Jan. 14. Antigua.

Jan. 15-Feb. 3. Pantaleon (1700 feet), Pacifie slope.
‘Tierra caliente. Sugar-cane fields and second
growth (rastrojo). ;

1881. Feb. 4-14. Mirandilla (1700 feet). Similar ground.

Feb. 15. Escuintla.

Feb. 16-18. Torola (1000 feet). Scattered patches of
forest.

Feb. 19-28. Paso Antonio (400 feet). Open savannas,
with scattered Crescentia and other trees, near Pacific,
above Istapa. Lagoons here adjacent to the Rio
Michotoya drained and used for pasturing the hungry
cattle in dry season. Silurians and other fish
captured in large numbers by the natives when the
water becomes low in the lagoons, and iguanas also
sought after, for food. Culicide and ticks very
troublesome. Some new Dytiscide and other aquatic
insects taken.

March 1-3. ‘Torola.

March 4. Savana Grande, near the Rio Maria Linda
(about 1150 feet). Scrubby woods, pasturage, and
cultivated ground. Passed through village of
Guanagazapa.

March 5. La Gavita (La Gavia) (1700 feet). Similar
ground. ;

March 6, 7. Brito.

March 8-16. Torola. District swarming with ticks
in dry season, and extremely unproductive, entomo-
logically, like the rest of the ‘tierra caliente’ of the
Pacific slope, during this period.

March 17-April 38. Guatemala city.

April 4,5. Escuintla.

April 6, 7. San Joséde Guatemala. Left by coasting
steamer on 7th en route for Panama.

April 9. La Union, Salvador. Landed for a few
hours.

April 10. Corinto, Nicaragua. Landed for a few
hours.

April 12. Punta Arenas, Costa Rica. Landed for a
few hours.

April 17-20. Panama city. Left on 20th by small
steamer for Chiriqui; the ‘“ port” reached by a
tortuous passage through mangrove-swamps, navi-
gable at high-water.

April 22-380. David. Principal town of Chiriqui.
Open savannas, with scattered, leathery-leaved, deci-
duous trees. Denser growth by river-side and
on hills adjacent, the latter productive, entomo-
loyically.

May 1-27. Finca Nance Bonito (about 2860 feet).
Southern slope of the Volcan de Chiriqui. Plenty
of forest, cleared in places for coffee-plantations, a
fine palm locally abundant beneath the larger forest-
trees, at about 2000 feet, just above the limit of the
savannas. Conifere altogether wanting in the district,
probably not reaching south of Nicaragua. Two
species of monkeys (Cebus hypoleucus and an Ateles,
almost certainly geoffroyz) seen. Productive locality.

1881.

1882.

ITINERARY.

May 28—June 25. Las Potrerillas (El Banco) (about
2500 feet). Similar ground, not far from Nance
Bonito.

June 26-July 13. David.

July 14-Aug. 1. Finca La Elvira. Similar to Nance
Bonito.

Aug. 2-8. Las Potrerillas.

Aug. 9-11. Boquete (3550 feet). South-eastern slope
of the Volcan de Chiriqui. .

Aug. 12-20. La Caldera (1500 feet). Savanna region,
with scattered trees. On old route from David to
Atlantic coast, practical on foot only, but rarely
used, all provisions having to be carried.

Aug. 21-Sept. 27. Las Potrerillas.

Sept. 28-Oct. 20. David.

Oct. 21-Nov. 28. Bugaba (1000 feet). Fine forests
here *, extending into Costa Rica, eastward and south-
ward savannas, with scattered trees. Rio Ascaria and
Rio Chiriqui Viejo descending through forest-country.
Sugar-cane and coffee cultivated to a small extent.
The place of residence of the brothers Trétsch and
other collectors formerly employed by Staudinger,
the well-known Lepidopterist.

Nov. 29. Jugales. Resting-place on way up to the
Potrero del Volean.

Nov. 30-Dec. 10. Potrero del Volcan (4000 feet).
Road from about 1000 feet above Bugaba rapidly
ascending through dense forest. Savannas up to
6000 feet, to the precipitous western slope of the
Volcan de Chiriqui (about 11,000 feet), and to the
abruptly descending banks of the Rio Chiriqui Viejo.
Dense forests westward and southward, in which
peccary are not rare, and jaguars, &e., occur. Large
numbers of cattle for the Panama market pastured on
these savannas. Monkeys (Ateles sp.) and peccary
(Dicotyles labiatus) sometimes shut for food by the
‘vaqueros. Uninhabited district.

Dec. 11,12. Bugaba.

Dee. 13-26. David.

Dee. 27-31.

Jan. 1-Feb. 2. } Bugaba.

Feb. 3-5. Camaron (1750 feet). Milpas (maize-fields)
in forest-clearings, on Costa Rica road.

Feb. 6—March 15. Bugaba.

March 16-21. Las Potrerillas. Country in a smoky

haze at this season, due to the general burning of the

undergrowth. Fires dangerous to the thatched
houses, owing to the strong ‘ northers.’

March 22—April 19. Bugaba.

April 11-13. David.

April 14-18. Bugaba.

April 19-29. Potrero del Volcan. Second visit, more

* Probably all destroyed by this time.

t

D3

productive than first. Many large Lamellicorn-
beetles (Plusiotis, various Dynastids, &c.) found
floating in the water of the large shallow lagoons in
the forest, on the banks of which numerous trees
frequented by the insects were in flower. The short-
tailed ‘quezal’ (Pharomacrus costaricensis) seen at
about 5500 feet.

1882. April 30-May 30. Bugaba. A very good locality at

this season, the commencement of the rains. New
clearings in forest—made to plant rice, maize, or
tobacco—productive in Coleoptera, Rhynchota, &c.
A large number of interesting Lepidoptera Rhopalo-
cera occur in the district, including Morpho eypris
and other species of the genus. A marmoset, Chiyso-
thrive oerstedi, occasionally seen in numbers on the
outer limits of the forest ; also an occasional peccary
(Dicotyles tapaju).

May 30-June 5. Las Potrerillas.

June 6-8. Slope of the Volcan de Chiriqui (5500 feet).
Staying in hut erected by Swiss orchid collectors on
the very steep forest-clad slope. Ascended to 8000
feet, by tracks made by tapirs through the dense
undergrowth of bamboo, &e., to summit of ridge.
Further progress stopped by an immense precipitous
ravine, beyond which the upper part of the volcano
could be clearly seen through the trees. The vege-
tation on the summit consisted apparently of low
bushes (? Vaccinium). No water could be found near
hut and a longer stay not possible. Some interesting
new Lampyridex, Hispid, and other Coleoptera found
in these places.

June 9-12. Las Potrerillas.

June 13, 14. Finca Nance Bonito.

June 15-July 38. Bugaba.

July 4. Jugales.

July 5-7. Potrero del Volcan.

July 18-Aug. 15. Bugaba.

Aug. 16-19. David.

Aug. 20-28. Bugaba.

Aug. 29, 30. Divala (Vivala) (350 feet), Savannas,
covered in places with a short prickly Mimosa, the
Rio Chiriqui Viejo adjacent. Tropical forest, with
many lofty palms, westward. A ‘road’ into Costa
Rica passes this place, not many miles from the
Pacific. Eastward, on the savanna, is the old
capital of the district, Alanje. Abandoned saw-mill
noticed on one of the streains.

Aug. 31. Mosque (700 feet).

Sept. 1-Nov. 30. Bugaba.

Dec. 1-3. David.

Dec. 4-27. Bugaba. Garrapatas (ticks) swarming in
the bush during dry season, necessitating a frequent
change of clothing. Boas occasionally met with.
Insects scarce, except near water, butterflies mostly

1883,

2, Dec. 28-30.

‘b4
worn. Many species of latter found about the
scattered pools of the nearly dry Rio Ascaria at this
season, iostly Nymphalids and Hesperids. Culicids
not very troublesome.

David.

Dec. 31. Chorcha (300 feet). Dense forest, descending
down to the mangrove-swamps of the coast, inter-
rupting the continuity of the large savannas border-
ing the Pacific east of David, said to be infested with
snakes.

Jan. 1. San Lorenzo (200 feet).
here and there.

Jan. 2, 3. Los Remedios.
through mangrove-swamps. Savannas inland.

Jan. 4-10. Tolé (1150 feet). Similar country. Ex-
cursions made from here to various places in Cordil-
lera. Cattle-breeding the principal business here.

Jan. 11. Cerro Algodon (2000 feet).

Jan. 12-17. Petia Blanca (3000 feet). Very broken
open country in vicinity of the Pena (Bluff), Slopes
of the mountains covered with forest. Ascended to
about 5500 feet. Indians from distant places on
the Atlantic (northern) slope assemble in an unin-
habited spot in the Cordillera near here annually.
Dancing in the open air is kept up for two or three
days, till most of the men and women become hope-
lessly drunk from the aguardiente supplied by
travelling traders. The adults seen at one of these
functions were tattooed with blue and red pigment.

Jan. 18. Cerro Algodon.

Jan. 19-22. Tolé. Nearly the whole of my money
stolen from saddle-bags at this place, a return to
David therefore imperative. Savannas east of Tole,
on the Santiago de Veraguas road, visited, but found
to be unproductive at this (dry) season. Several
interesting Malacoderm-beetles (Astylus, Lycostomus,
&c.) were, however, taken from flowers. Very little
cultivation seen.

Savannas with woods

A small port, reached

Jan. 23-28. Nancito (800 feet).
Jan. 29. Los Remedios.

ITINERARY.

| 1883.

Janu. 30-Feb. 3. San Feliz (650 feet). Savannas, with
patches of wood.

Feb. 4,5. Lalsleta. Adjacent to the Rio Fonseca. A
‘ Howler’ (Mycetes palliatus) seen in the trees along
the streams.

Feb. 6, 7.
Chorcha.

Feb. 8-10. David.

Feb. 11-March 3. Bugaba.

March 4-9. David.

Mareh 10,11. Bugaba.

March 12,13. David.

March 14. Boquita, on way down to coast.

March 15-17. Boca Chica, near coast.

March 18-24. En route to Panama in small coasting-
vessel carrying various passengers, who were accom-
modated in the open air on planks above the cargo
of pigs. Punta Mala passed with difficulty.

March 25, 26. Panama.

March 27. Colon (Aspinwall). Many butterflies seen
from train on way across isthmus, but next to none
found at Colon.

March 28-April 2. Panama.

April 3-27. San Miguel, King Island (Isla del Rey),
PearlIslands. Scrubby woods much cleared in places
to plant yams, &c. Tangled jungle, with some large
trees, in southern portion of island. Coast fringed
with mangroves and coco palms. Darien coast and
mountains visible from San Miguel. A few interest-
ing insects obtained. Unhealthy place. Too unwell
to do much collecting. No horses or cattle. Tracks
available along beach at low water.

April 28, 29. On way back to Panama in small boat.

April 30. Panama.

May 1-17. Taboga Island, Bay of Panama. Rocky
ground, ascending to about 800 feet, with small
streams. Pine-apples grown on the slopes, often
stolen by sailors. Many butterflies and other insects
occur on the island.

May 18-21. Panama.

Cafia Fistula. Near the Montana de

rm
Or
Cu
Li

LIST OF COMPLETED VOLUMES.

ZooLoey, Botany, AND ARCHAOLOGY.

Tue 215 Parts of Zoology, 25 of Botany, and 17 of Archaeology are divided into
63 Volumes, of which a complete list, with their contents, is given in tabular form

on pp. 55, 86. The analysis of the contents of each of these volumes supplies the
names of the contributors and other particulars, and, in the case of the Zoology,

a brief summary of the author's views on the nature of the Fauna, as stated in
their Introduction. The 215* Parts of Zoology form 52 volumes—one devoted
to Mammalia, four to Aves, one to Reptilia and Batrachia, one to Pisces, one to
Mollusca, four to Arachnida, one to Chilopoda and Diplopoda, and thirty-eight to
Insecta. The Crustacea, Protobracheata, Vermidea, &c., have not been studied,
mainly for lack of material. Amongst the Insecta, too, no worker has been found for
certain groups of Hymenoptera, Diptera, Rhynchota-Homoptera, and Neuroptera, and
these omissions are specially noted in the analysis of the volumes dealing with the
Orders in question. The 25 Parts of Botany form five volumes—four of text and one
of plates. The 17 Parts of Archeology form four volumes of text, together binding
into one volume, with an additional common titlepage, four volumes of folio plates,
and an Appendix (text only), the whole subject thus extending to six volumes—two

of text (quarto) and four of plates (folio).

* Part 211 was issued in two sections: ‘ A” in Dec. 1911, “ B” in May 1912.
Part 212 » ” » +: “A” in Feb, 1913,“ B” in April 1914,

[ 56 |

ANALYSIS OF CONTENTS OF EACH VOLUME.

1. The ‘Introductory Volume’ includes the general preface to the whole work,
Zoology, Botany, and Archeology, a complete list of the subjects contained in each
Volume, general statistics, maps, &c. The ‘Contents’ of this Volume, which forms
the first of the entire series, are given in detail on p. li.

ZOOLOGY.
2. Mammatia: by E. R. Alston, with an Introduction by Dr. P. L. Sclater.

The author of this Volume died on March 7th, 1881, before the enumeration of the
species was concluded. The MSS., however, of the Supplement (pp. 203-212) was
left by Mr. Alston in an almost complete state, and was finished by Mr. O. ‘Thomas in
1881. Vhe Tables (I-VIIIL.), printed in the Introduction, were drawn up at our
request by the author shortly before his death, and Dr. P. L. Sclater gives an
analysis of them in his Introductory remarks on the subject (pp. x—xix), published in
1882. The total number of species enumerated is 181, of which a complete list is
given in the Introduction (pp. iv-ix). These fall into five categories: (i.) Nearctic, 48 ;
(ii.) Neotropical, 65; (ili.) Neogean, 17; (iv.) Autochthonous, 47; and (v.) Intro-
duced, 4 (AZures). The eight Tables drawn up by the author show: I. General dis-
tribution; II. Nearctic species (17) found in the Northern States of Mexico, but not
recorded from south of 25° N. lat.; iI. Nearctic species (18) found in Central and
South Mexico, but not recorded from south of the Isthmus of Tehuantepec;
IV. Nearctic species (8) extending to Guatemala and Honduras, but not recorded
from south of Nicaragua; V. Nearctic species (5) extending to Costa Rica and
Panama, but not recorded from south of the Isthmus of Panama; VI. Species (17)
common to the Nearctic and Neotropical Regions; WII. Species (47) peculiar to
Central America, or not yet ascertained to occur elsewhere; VIII. Distribution of the
Neotropical genera. Dr. Sclater (p. xiv) summarizes the analysis of the Central
American Mammal fauna as follows :—“It may fairly be said that (excluding the
introduced Mures) at least 100 of the constituent species are essentially Neotropicalin
their character or have Neotropical! affinities, while of the remainder not above 60 can
be said to be decidedly Nearctic. There is therefore no doubt that the Central
American isthmus, at any rate as far north as Tehuantepec, should be assigned to
the Neotropical Region, of which it should be deemed to constitute a distinct province,
characterized (1) by the incursion of a considerable number of Nearctic forms,
especially in the northern districts, (2) by the presence of a certain number of peculiar

ZOOLOGY. 57

species of Neotropical genera, and (3) as being the focus of the families Procyonide
and Geomyide, two well marked groups of Mammals which have extended alike
into the Nearctic and Neotropical Regions.”

The twenty-two coloured plates include figures of thirty-four species, a list of which
is given on p. Xx.

3-6. Aves: by O. Salvin and F. D. Godman: Vols. I-III. (text), III. completed
with the assistance of Dr. R. Bowdler Sharpe and Mr. Ogilvie-Grant ;
IV. (plates).

Three Volumes are required for the enumeration and description of the 1413 species
of Aves belonging to the Central American fauna, and a fourth for the 84 plates.
Vol. I., published in 1879-1887, gives an account of a portion of the Passeres, the
families Turdide to Alaudide; and, on the conclusion of the work, in 1904, an
Introduction to the whole subject was issued, with Tables (pp. xi-xxxvili) showing the
geographical distribution of the families and species represented in Mexico and Central
America. Vol. II., published in 1888-1897 *, includes the continuation of the Passeres
and the whole of the Macrochires, Pici, Coccyges, and Psittaci. Vol. III., published
in 1897-1904, includes the Striges, Accipitres, Steganopodes, Herodiones, Phoenicopteri,
Anseres, Columbe, Galline, Geranomorphe, Limicole, Gavi, Tubinares, Pygopodes,
Alce, and Crypturi. Vol. IV. contains the whole of the Plates and a complete list of
the 149 species figured. Salvin’s long continued. ill health, and sudden death in 1898,
retarded the conclusion of the Third Volume, and this was subsequently finished with
the assistance of Dr. Sharpe and Mr. Ogilvie-Grant. For this reason, too, all idea of
a Supplement was abandoned, notwithstanding the large amount of additional material
which had come to hand during the progress of the work. The additions, however,
were mainly amongst the Passeres, completed in 1892. The Introduction to Vol. 1.
(1904) contains an account of the authors’ various expeditions to Central America,
the sources from whence their material was obtained, &c., and the following par-
ticulars as to the nature of the Bird-fauna of the region :—To summarize the results,
the Avifauna of Central America may be described as essentially Neotropical, with
certain peculiar forms restricted to it. The fifteen families represented are all rich in
endemic forms, and the families themselves are almost all tropical. On the other
hand, a large number of species belonging to the more widely distributed genera find
their winter home in Mexico and Central America, or further south, returning to
breed in the Nearctic Region, even Humming-birds and others wandering far north at
this season. The data is insufficient to show the lines of migration of all the species.
Some, no doubt, travel southward from the United States to the mainland of South
America by way of the Caribbean Sea or the West Indian Islands, perhaps just

* The permanent Titlepage and ‘ Contents’ were issued in 1904.

BIOL. CENTR.-AMER., Introd. Vol., January 1915. I

58 CONTENTS OF EACH VOLUME.

touching the eastern portion of our region en route; others probably find their way
down to the central tablelands, and a few western species, again, pass over the low
lands of the Pacific coast.

Nearly half (636) of the 1413 species enumerated are treated as endemic, fifteen
of the families—Trochilide, Fringillide, Tanagride, Formicariide, Dendrocolaptide,
Troglodytide, Turdide, Psittacide, Phasianide, Peristeride, Cotingide, Trogonide,
Cracide, Tinamide, and Rhamphastide—having many peculiar forms. Compared
with America north of Mexico, the fauna of the region here dealt with is particularly
rich, the numbers being, North America (1895) 768 species as against Central
America 1413; while India (with Ceylon and Burma), with nearly double the area, has
(1898) only 1626 species. All, or nearly all, the new species were described first in
the ‘Ibis’ or ‘P. Z. S.’ by Salvin.

Birds, from their power of flight and the habit of migration common to a large
number of them, are much more easily distributed than most other vertebrates, and,
therefore, do not throw the same light on the subject of geographical distribution
as in the case of more sedentary animals. This must, nevertheless, be applied in
a general sense, for many of the species are extraordinarily local.

In dealing with the Aves the limits of the region have been extended to include
the Revillagigedo Islands on the Pacific side, on account of the numerous sea-birds
inhabiting them; the Island of Old Providence on the Atlantic side, which has
a humming-bird peculiar to it; and some places on the Isthmus of Darien.

The eighty-four coloured plates illustrate 149 species.

7. Reprivia and Barracuta: by Dr. A. Giinther.

The author, in his Introduction published in 1902, summarizes his remarks on
geographical distribution as follows:—The general features of the Reptilian and
Amphibian Faunas of the area under investigation have been satisfactorily ascertained.
Forming the connecting link between the two Neogean regions, Central America
possesses a Reptilian and Batrachian Fauna with the various constituent elements so
mixed that, if only certain families or genera were taken into consideration, almost
every district of this area could be associated with either the North- or South-American
region. ‘The tropical Fauna, as we proceed from lower to higher latitudes, gradually
changes or is replaced by that of the temperate region; but this change is not
uniform throughout the breadth of the land, and the two faunas frequently overlap
in deep and manifold indentations. ‘Tropical types are found to preponderate in
the low lands of the Atlantic side, which expand into the broad Yucatan peninsula,
and on the humid slopes of moderate elevation; some extend to, and even reach
northwards of, the Rio Grande. On the Western side they are found in similar
localities, but in a narrower belt, along the Pacific coast. On the other hand,

ZOOLOGY. 59

numerous types of the southern North American Fauna are spread over Northern
Mexico, extending along the Central American plateau to the extreme limits of our
area, and even beyond. This southward extension of northern types is due partly
to the identical physical conditions of the arid tableland of Sonora and Chihuahua,
which is merely a continuation of that of Arizona and New Mexico, and partly to the
great altitude and temperate climate of the Central American plateau. Thus, a
boundary line between the North and South American regions cannot be drawn:
Central America forms a transition-tract unlike any other part of the world, showing
the most extraordinary diversity of climatic, physical, and meteoric conditions
within comparatively small areas, favouring the evolution of a great variety of types
of genera and species, and influencing the dispersal of immigrants from the North
and South.

The range of the 695 species enumerated is shown in the Table appended to the
Introduction (pp. x—xvil).

Since the conclusion of Dr. Giinther’s work, Dr. Hans Gadow has twice visited
Mexico (1902 and 1904), mainly to study the distribution, &c., of the Amphibians and
Reptiles. He collected specimens of 135 species. The following papers have been
written by him :—

(1) ‘Evolution of the Colour-pattern and Orthogenetic Variation in certain
Mexican Species of Lizards, with Adaptation to their Surroundings ” [ Proc.
Roy. Soc. Lond. Ixxii. pp. 109-125, pls. iii.—v. (1903) ].

(2) “The Mexican Axolotl” [‘ Nature,’ lxvii. pp. 330-332 (1903) ].

(3) “The Distribution of the Mexican Amphibians and Reptiles” [P. Z. S. 1905,
ii. pp. 191-245].

(4) “ A Contribution to the Study of Evolution based upon the Mexican Species of
Cnemidophorus” (P. Z. 8. 1906, i. pp. 277-375, pl. xx.).

(5) ‘Through Southern Mexico.’ London, 1908.

(6) ‘“‘ Geographical Distribution of Animals” [‘ Darwin and Modern Science,’ pp. 319-
336 (1909).

(7) “The Effect of Altitude upon the Distribution of Mexican Amphibians and
Reptiles” [Jahrb. Jena Abt. f. Syst. xxix. 1910, pp. 689-714].

He also published an excellent account of the portions of Mexico visited during his
travels, and we give elsewhere a copious extract from one of his papers.

During recent years attention has been called to various Reptilia, Batrachians,
Arachnids, Coleoptera, &c., living in the Bromeliads on the branches of trees.
Specimens of Spelerpes variegatus and Hyla godmani were obtained from these plants
by one of our collectors, Mateo Trujillo, in Mexico, and other species have since been

captured by Dr. Gadow.
. 12

60 CONTENTS OF EACH VOLUME.

8, Pisces: by C. 'T. Reaan.

The Introductory remarks to this subject (published in 1908) are arranged by the
author under five headings -—(1) Principal Faunal Works on the Fresh-water Fishes
of Mexico and Central America; (2) Principal Collections described in this Work ;
(3) Classification ; (4) Geographical Distribution, illustrated by two separate maps;
(5) The Shore-Fishes of the Atlantic and Pacific Coasts of Mexico and Central
America. The geographical distribution is discussed at length under the headings to
the separate families, ten of which are represented in the region, that of the Cichlide,
Percide, Characinide, Catostomine, and Cyprinine being illustrated by shaded maps in
the text. The author considers that the Nearctic and Neotropical Regions are quite
distinct, and to explain the distribution of fresh-water fishes he says that the volcanic
chain of mountains which stretches across Mexico from Colima nearly to Vera Cruz
may be taken as the boundary. The Nearctic Region he divides into three subregions,
one of which, the “Lerma,” lies wholly within Mexican territory. The Neotropical
Region is also divided into three subregions, the Central-American subregion again
being separated into four “ provinces,” Balsas, Guatemalan, San Juan, and Isthmian ;
the limits of these are shown on Map 2.

Mr. Regan says that the Cichlide, which form a very large family of fresh-water
fishes, is the dominant perciform group in Tropical America and Africa. In America
the Cichlide extend from Texas to Argentina, and comprise about 150 species. Africa
appears to be somewhat richer in both genera and species, whilst three representatives
occur in India and Ceylon. A mapis given in the text to show this distribution. He
states that the Mexican and Central American Cichlids must have originated in the
Southern Continent.

‘The Percide are fresh-water fish inhabiting Europe, Northern Asia, North America
east of the Rocky Mountains, and Northern Mexico. Fossil Percide are found in
the Eocene deposits of Wyoming.

The distribution of the Characinide, also fresh-water fish, is equally interesting.
They extend from Northern Mexico over all the South-American continent, except
the extreme south, and are found also in Africa, but not in Madagascar.

These two families and the Lepidosirenide support the theory that a land
connection between South America and Africa may have persisted until the beginning
of the Tertiary period.

The Catostomine extend from Guatemala all over North America and into Eastern
Asia.

The Cyprinine are found from Canada southward to the Lerma Valley in Mexico
and throughout Africa and Eurasia, except in the extreme north. ‘They abound
in Borneo, but are absent from Celebes.

Amongst the Shore-Fishes entering fresh water, eighteen Pacific coast species are
specially noted as having a closely related representative on the Atlantic.

ZOOLOGY. 61

Mr. Regan concludes his remarks on these fishes as follows :—‘ The fact that in so
many cases species may be paired is more in harmony with the view that there has
been a gradual modification during isolation than with the supposition that a mutant
has arisen which has replaced the parent form.”

The twenty-six plates include figures of seventy-eight of the 415 species enumerated.
The Maps inserted in the Introduction showing the distribution of certain families are
worthy of attention.

9. TERRESTRIAL and FiuviatineE Mo.uusca: by E. v. Martens.

In this volume the author summarizes the characters and distribution of the species
in common Tables, instead of giving separate descriptions. The geographical distri-
bution is given at great length in the ‘ Introduction’ (published in 1901), accompanied
by separate Tables of the Terrestrial and Freshwater genera. The difference between
the fauna of the Pacific and Atlantic slopes is noticed, but is stated to be not very
important as regards the land shells. Central Guatemala (Baja Vera Paz), however,
is said to be occupied chiefly by metamorphic rocks, while North Guatemala (Alta
Vera Paz) is mostly of limestone formation: this difference accounts for the greater
richness of land shells in the latter province. One peculiarity of the fauna of the
Pacific slope is worthy of note, viz. the occurrence of large sized Otostomi and Bulimult
in Western Mexico related to various Andean forms, the distribution being somewhat
analogous to that of the majority of the Cactacee. The submarine Mollusca of the
eastern and western shores are said to be distinct, more so than some truly marine
shells, but there are some remarkably analogous forms among them.

Of the forty-four plates belonging to this subject, the first twenty-eight are
coloured.

[The Crustacea have not been studied.
The FresHwaTerR Matacostraca were undertaken by Prof. T. H. Huxley,
but his contribution was never finished. |

10. ARACHNIDA ARANEIDEA. Vol. I.: by the Rev. O. Pickard-Cambridge.

This volume contains descriptions and figures of 417 new species of Araneidea, and
a list of species identified by the author. The new forms are illustrated on the thirty-
nine coloured plates.

11. ARAcHNIDA ARANEIDEA and Opruiones. Vol. II.: by F. O. Pickard-Cambridge.

Vol. II. of this subject gives the systematic arrangement of the whole of the species
of Arancidea described in Vol. I. and the large number added in Vol. II. A synopsis
of the Families is given on pp. 541-544. The ‘Opiliones,’ or ‘ Harvestmen,’ are
dealt with on pp. 546-585, and illustrated by three plates, LIL—LIV. ‘The author is
unable to say very much about the general distribution of the Araneidea, the material

62 CONTENTS OF EACH VOLUME.

at hand being insufficient for the purpose. They are said to date back at least as
far as the Carboniferous period, when they were represented by forms of which the
two species of Liphistius are the sole survivors. The earliest known Arachno-
morphid spiders in any way resembling those living belong to the Oligocene times.
At that period there existed many forms, very similar to those occurring at the present
time, whose remains have been found in amber washed up on the shores of the Baltic
Sea. The Opiliones are stated to have preceded, during the Carboniferous epoch, the
air-breathing scorpion, Anthrascorpius, and their specialization probably began during
the still earlier Silurian times, since in that period there existed an Arachnid which
is a true scorpion in every sense, except that it had apparently no trace of air-breathing
lung sacs. The ‘ Harvestmen’ make no web for the ensnaring of their prey, and the
females, after depositing their eggs, take no further interest in their offspring. In the
two volumes 1181 species are enumerated, this number including the 422 described
as new in Vol. II. ‘The Opiliones number 70 species, 58 of which are described as
new, with 11 new genera. The ninety-three plates (39 in Vol. I. and 64 in Vol. II.)
include figures of 981 species.

12. ARACHNIDA ScorPIONES, PEeprpatri, and Souiruc#: by R. I. Pocock.

Very little material was available for the study of these Arachnids, sixty-nine species
only being enumerated for the three groups. The twelve uncoloured plates include
figures of thirty-seven species. The Scorpiones are represented by three families, the
Pedipalpi by two, and the Solifuge by a single family.

In this Volume the author gives the geographical distribution under the heading
for each genus, and no general ‘ Introduction’ to the whole subject was prepared, for

want of data.

13, ARACHNIDA ACARIDEA: by O. Stoll.

‘The material for this subject was obtained almost entirely by the author during a
residence of nearly five years in Guatemala. He made the drawings on the spot, but
unfortunately he had no modern literature on Acarids with him, and his microscope
was anything but satisfactory. Dr. Stoll’s work, therefore, must be treated as a
contribution to the fauna of Guatemala, rather than as an enumeration of the
Acaridea inhabiting Central America. The types of the species described remained
in his possession. In his ‘Introduction’ (published in 1893), p. vii, he states that it
is remarkable that not one of the forms described represents a generic type entirely
new or peculiar to the region. Doubtless a great deal remains to be done in the
way of collecting before we shall have any true idea of the Acarid-fauna of Central
America.

The twenty-one coloured plates illustrate 43 species.

ZOOLOGY. 63

14. CurLopopa and DipLopopa: by R. I. Pocock.

The dates of publication of this Volume are, Chilopoda 1895-1896, Diplopoda
1903-1910. For the two groups 255 species are enumerated, of which 106 are
described as new, the Chilopoda numbering 53 (19 new) and the Diplopoda 202
(87 new) respectively. The author, for want of sufficient data, does not give any ©
particulars as to general distribution, beyond that mentioned under each genus or
species in the text. ‘The three plates belonging to the Chilopoda are partly coloured,
the twelve others referring to the Diplopoda are uncoloured.

[The Prororracuzata (Peripatus, &c.) have not been studied.
One species at least has been recorded from Nicaragua. |

15. Cotnoptera. Vol. I. part 1: by H. W. Bates: Cicindelide and Carabide.

The author, who had previously studied the insect fauna of the Amazons during his
long residence in that region, remarks, in his ‘ Introduction’ (published in 1884), on
the Central-American fauna of these two families as follows :—‘‘ The number of species
(1086), belonging to 154 genera, is greater than the apparent poverty of tropical regions
in Carabide would have led us to expect. The tolerably well-worked valley of the
Amazons, although rich in species of genera belonging to alluvial plains, and in
arboreal forms, contains only 576 species belonging to 124 genera; and the fauna of
such tropical regions as the Malay archipelago is still poorer. The reason for the
comparative paucity of Carabide has been supposed, apparently on good grounds, to
be that their place, as predaceous terrestrial insects, is to a great extent occupied by
the ubiquitous ants. The undoubted fact that purely epigzeous Carabidee, except
marsh species, are scarce in the Tropics, especially near the Equator and in the low-
lands, and that arboreal or climbing forms alone are numerous and varied, affords
support to this hypothesis. ‘The essentially Neotropical character of the Central-
American fauna is generally admitted, and is strikingly confirmed by the Cicindelidze
and Carabide. But with regard to the northern limits of the fauna, and especially
the extent to which Nearctic and North-temperate forms have penetrated the region
from north to south, these are points not yet settled. Wallace included, or seemed
inclined to include, the whole of the central highlands of Mexico and Guatemala in the
Nearctic province, which must mean that the North-temperate American forms are
there in the majority. The two families of Coleoptera we are dealing with do not
support this conclusion. The Nearctic forms are comparatively few, and in the ‘ tierra
templada’ are far outnumbered by tropical genera. The northern limit of the Central
American fauna appears to be—on the highlands, if not also on the maritime lowlands
east and west—a little south of the political frontier of Mexico. Does the Ceutral
American fauna constitute one homogeneous province, or is it divisible into two
subprovinces, as Salvin (‘ Ibis,’ 1866, p. 202) has shown to be probably the case with

64 CONTENTS OF EACH VOLUME.

the birds, many genera of which are represented by distinct species on each side of a
line which he is inclined to place north of the Nicaraguan lakes and their outfall, the
Rio San Juan ?”

‘The thirteen coloured plates include figures of 324 species.

Mr. Bates subsequently wrote two papers on the Mexican materia] received from our
collectors afier the present Volume was closed :—

(1) ‘ Additions to the Cicindelide Fauna of Mexico’ (Trans. Ent. Soc. Lond.
1890, pp. 4938-510, pl. 16).

(2) ‘Additions to the Carabideous Fauna of Mexico’ (op. cit. 1891, pp. 223-
277, pls. 18, 14).

16. Cotzoprera. Vol. I. part 2: by D. Sharp: Haliplide, Dytiscide, Gyrinide,
Hydrophilide, Heteroceride, Parnide, Georisside, Cyathoceride, and
Staphylinide.

For these nine families of Coleoptera 1790 species are enumerated, 1405 of which
belong to Staphylinide, 168 to Dytiscide, and 141 to Hydrophilide, the others being
of limited extent. The author, in his ‘Introduction’ (published in 1887), states that
the water beetles (Dytiscide) are apparently subject to different laws of distribution
from other Coleoptera, illustrating in this respect what he believes to be the case
with aquatic organisms generally, viz. diminished endemicity, and therefore but little
adapted for consideration in questions of zoo-geography. The study of the exotic
Staphylinide appears to be practically in its infancy, and but little can be done in the
way of comparison with the fauna of other tropical regions. In the case of genera
with a large number of species and a wide distribution, the Central American
Staphylinids are stated to be much more nearly allied to those of South than to
those of North America. Dr. Sharp is therefore inclined to agree with the opinion
expressed by Bates that the fauna of the region under investigation is essentially
Neotropical. The Staphylinide first brought from the tropics included an undue
proportion of comparatively large, bizarre, or brilliantly coloured forms, but it is clear
that this was chiefly due to imperfect collecting. Indeed, one of the striking facts in
connection with the material brought together by the Editors is the very large number
of minute and obscure insects, so that it can scarcely be said that the Central American
Staphylinide are, on the average, larger or more brilliant than those of Europe.

Of the nineteen coloured plates, fourteen are devoted to Staphylinide, 445 species
being figured altogether.

17. Cotroptera. Vol. II. part 1: by Dr. Sharp, A. Matthews, and G. Lewis:
Pselaphide—Byrrhide.

The twenty-two families enumerated in this volume are collectively termed
Clavicornia by various authors, and in recent catalogues of Coleoptera the Erotylide,

ZOOLOGY. . 65

Endomychide, and Coccinellide (these three families being here placed at the end of
the Coleopterous series in Vol. VII.) are treated as belonging to the same division.
The Silphide, Corylophide, Trichopterygide, Spheriide, and Scaphidiide were
worked out by Mr. Matthews, the Histeride by Mr. G. Lewis, and the remaining
sixteen families by Dr. Sharp, the last named author also contributing a list of the
Rhipidandri—a small group of somewhat uncertain position, but really belonging to
the Tenebrionid-series near Boletophagini. The total number of species is 1629, of
which 996 are described as new. ‘The dates of publication of the various subjects are:
Pselaphide and Scydmenide, 1887; Silphide—Scaphidiide, 1887-1888; Histeride,
1888; Phalacride—Byrrhide, 1888-1905; Rhipidandri, 1905. The ‘ Introduction’
to this Volume was simply an editorial note; but in his remarks on the Histeride
(p. 182) Mr. Lewis states that the chief interest of this series of insects centres
in the species which feed on the wood-boring Coleoptera of other families. ‘The
nineteen uncoloured plates include, it is believed, some of the finest lithographic
illustrations of beetles that have as yet been published, those devoted to the Colydiide
(plates xiv. and xv.) being particularly excellent. The artist, Baron Max Schlereth,
was unfortunately unable to draw the whole number.

18. CoLeortera. Vol. IL. part 2: by H. W. Bates: Pectinicornia and Lamellicornia.

The total number of species for these two impcrtant families is 1100—72 Pectini-
cornia and 1028 Lamellicornia. ‘The author, in his ‘ Introduction’ (published in
1890), remarks that the Pectinicorn-fauna is exceedingly poor in the chief family of
the tribe, viz., the Lucanide; but, on the other hand, it is unusually rich in the more
aberrant family, the Passalide. A comparison with such allied faunas from other
parts of the world, which have been sufficiently worked out to give approximately
accurate results, seems to show that the poverty in Lucanide arises from Central
America lying too far south to have been reached by many species of Old-World genera,
‘and too far north for the genera characteristic of South Brazil, Chile, and the Andes.
The conditions seem, however, to be very favourable to the Passalidz, which all pass
their earlier stages in rotting tree-trunks, reaching their highest development and
exhibiting more diversity of form here than in any other region. All the families
of Lamellicornia are well represented in Central America; they include 1028 species
belonging to 127 genera, but a comparison of the Lamellicorn-fauna with that of
other tropical regions of similar extent is impossible, as the necessary data do
not exist ina connected form. The actual number of species described up to 1890,
for the whole world, may be roughly estimated at 10,000. Of the twenty-four plates
illustrating the 492 species figured, all but one (Pectinicornia) are coloured.

Since this Volume was finished (in 1890) a good many species of Lamellicornia
have been added by various authors, these showy beetles being great favourites

BIOL. CENTR.-AMER., Introd. Vol., January 1915. K

66 CONTENTS OF EACH VOLUME.

with collectors, but the additions do not materially affect the conclusions arrived
at by Bates.

19. Cotzoprera. Vol. III. part 1: Serricornia: Buprestide by C. O. Waterhouse ;
Throscide and Eucnemide by G. H. Horn; Elateride *_Dascillide by
G. C. Champion.

For these families of the Serricorn-series, 1353 species are enumerated from Central
America, 805 of which are described as new. The Buprestide were published
sn 1882-1889: the Throscide and EKucnemide in 1890; the remaining families in .
1894-1897 ; and a short Appendix in 1897. The Buprestide, numbering 434 species
(exclusive of those mentioned in the Appendix), do not appear to be very much in
evidence, apart from the gigantic Huchroma, in the tropical forests of Mexico and
Central America, their place being to a great extent taken by the conspicuous Elaterid-
genera Chalcolepidius and Semiotus. The open parts of Mexico, however, have a rich
Buprestid-fauna, as shown by the number of species obtained by Hoge in his later
expeditions to that country ; these were enumerated in the ‘Supplement’ (published
in 1889). The distribution of Conifers, which do not extend south of Nicaragua,
probably affects the range of certain genera of this group. ‘The Elateride, numbering
531 species, have exceedingly few endemic genera, and it may be said in a general
way that the tropical forms are mostly confined to the forest regions, and that many
of those inhabiting the open country or higher ground are nearly allied, or actually
belong, to Nearctic genera. Pyrophorus has one species in North and sixteen in
Central America, and is essentially Neotropical. The Dascillide, with 130. species
and twenty genera, is one of the ‘neglected’ families of Coleoptera, and there are no
available data for comparison with other regions. The Eucnemide, of which a table
of the genera is given by Horn (pp. 211-213), are represented by 113, the Throscidee
by forty-four, the Cebrionide by twenty-nine, and the Rhipideceride by fourteen
species.

The twenty-seven plates, one of which shows the form of the terminal segment of
the males of the Buprestid-genus Pachyscelus, illustrate 648 species.

20. Conzoptera. Vol. III. part 2: by H. S. Gorham: Malacodermata.

This Volume includes the following nine families :—Lycide, Lampyride, Tele-
phoride, Lymexylonide, Melyridx, Cleride, Ptinide, Bostrychide, and Cioide. The
total number of species enumerated is 813; but as very many of our specimens added in
the ‘Supplement’ were not critically examined by Mr. Gorham, it is probable that the
actual number represented in the material obtained by our collectors is about 900.
The author, in his ‘Introduction’ (published in 1886), states that it is now seen that

* First undertaken by E. W. Janson,

ZOOLOGY. 67

the tropical portions of the earth are as rich, or richer, in these groups (he presumably
refers to Telephoride) than the cooler parts. ‘The Lycide, Telephoride, Lampyridae,
and Cleride clearly show a closer relationship between our fauna and _ that
of Tropical South America than with any they possess with that of America
north of Mexico—this being rendered even more apparent in such large genera as
Calopteron amongst the Lycids and Chauliognathus in the Telephorids. More striking,
however, is the marked peculiarity of the Central and South American genera as
compared with the faunas of other parts of the globe. The author remarks :—‘ If it
were true that similar conditions would produce similar forms, how is it that the
specialized genera of the Neotropical regions are so dissimilar from those of the
Asthiopian and Indo-Malayan? Of the few genera common to Central America and
to the Palearctic or Tropical regions of the Old World, there is not one which is not
of the rank of feebly differentiated forms, or persistent forms of an earlier stage of
development. The total absence of many widely distributed well marked forms cannot
be explained by any intrinsic dissimilarity in the conditions, which certainly do not
differ more than they do in the wide areas over which such genera are dispersed, but
rather by the isolation from these other areas, contact with which is only effected
through the northern parts of the globe. One further fact is noted, viz., that, to
whatever cause it may be due, where a genus is common to Central or South America
and other distant parts of the World, it is also the case that it is represented by a
species also identical or nearly related in the two districts. In this case its presence
must be due to recent transmission, through artificial means, or to the conditions
having long remained practically similar under which it exists in places so far apart.”

Thirteen coloured plates are issued in this Volume, 330 species being figured.

Some of the Clerids received by us since the conclusion of Mr. Gorham’s work have
been described by Herr 8. Schenkling in German publications, and a certain number
of Bostrychide have been dealt with or revised by M. P. Lesne. The Chauliognathine,
Malachiide, and Melyridee, and a portion of the Ptinide, have also been revised by
Mr. G. C. Champion (Trans. Ent. Soe. Lond. 1913, 1914).

21, 22. Cotzorrera. Vol. IV. parts 1 and 2: by G. C. Champion: Heteromera.

The fourteen families included in the Heteromerous-series are dealt with in two
volumes: (1) Tenebrionide, Cistelide, Othniide, Nilionide, and Monommide ;
(2) Lagriide, Melandryide, Pythide, Gtdemeride, Xylophilide, Anthicide, Mordel-
lide, Rhipidophoride, and Meloide. ‘The general ‘ Introduction’ (published in 1893),
which includes a Table showing the geographical distribution of the genera repre-
sented within the limits of Mexico and Central America, is given in part 1. The
total number of species enumerated is 1776, of which 1295 are treated as new.
Amongst the Tenebrionidee, the apterous terrestrial forms clearly belong to the

| K 2

68 CONTENTS OF EACH VOLUME.

Nearctic beetle fauna which extends down the central plateau to the Isthmus of
Tehuantepec (Asida) or to the Los Altes region of Guatemala (Zwodes), while the
winged forest forms are Neotropical. The winged genera inhabiting the open country
(Epitraqgus, Blapstinus, Xystropus, &c.) have a wide distribution, and such apterous
forms as appear to be insensible to drought and heat (e. g. Zopherus) range from
the Southern United States through our region to Colombia and Venezuela. The
humid forest regions of Central America possess a very rich Tenebrionid—fauna, as
so many species attack decaying trees, or the fungi growing upon them, even in
gloomy places; the Melandryide, however, are poorly represented, as in other
tropical regions. The Cistelidee, Xylophilide, and CEdemeride belong to the
‘neglected’ groups, not a single Central American species of any one of these families
having been described before the publication of these volumes. The Meloidze, on
the contrary, had been studied by various Coleopterists, especially by E. Dugés in
Mexico, and many of the species were previously described. ‘The Heteromera, as
stated in the Introduction, comprise a greater variety of forms than any of the other
main divisions of the Coleoptera, nearly all of which are reproduced here. Probably
no better case of mimicry or homochroism can be found amongst beetles than
that existing between the Tenebrionid genus Cuphotes and Cypherotylus (Erotylide)
and the Lagriid genus Uroplatopsis and Uroplata (Hispidee). Some of the smaller
subcortical Cucujids of the Clavicorn-series are so closely related to various Pythids—
from which they are only distinguishable by the number of joints in the hind tarsi of
the males—that it is probable that these latter will have to be removed eventually
from the heterogeneous series of families known collectively under the name
Heteromera.

The two volumes devoted to this group include forty-four coloured plates,
illustrating upwards of 1000 species.

23. CotzopreRA. Vol. IV. part 3: by D. Sharp and G. C. Champion: Curculionidee
(part).

The Rhynchophora, or weevils, include a larger number of species (3848) than any
of the other main divisions of the Coleoptera, and no fewer than five volumes of the
present series are devoted to their enumeration. In the First Volume (Part 3) 616
species of the following subfamilies of the Curculionide are dealt with: Attelabine,
Pterocoline, Allocorynine, Apionine, Thecesternine, and Otiorrhynchine. Dr. Sharp’s
contribution, pp. 1-177, was published (pp. 169-177 excepted) in 1889-1891; this
portion included the first five subfamilies and the apterous Otiorrhynchinx, and that
of Mr. Champion, dealing with the winged Otiorrhynchine, appeared in 1911. ‘The
Attelabine and Apionine, both numerous in species, do not differ greatly from the
representatives of these subiamilies in temperate northern regions. The Pterocoline,

ZOOLOGY. 69

Allocorynine, and Thecesternine, each including a few species only, are common to
North America and Mexico or Central America. The apterous Otiorrhynchids are
mostly restricted to the central plateau—Hupagoderes, Epicerus, and Epagriopsis
being the dominant genera in the highlands of Mexico,—while the winged forms
preponderate in the warmer forest regions to the east, west, and south, this
distribution being similar to that of the Tenebrionids. Pandeleteius and Pantomorus
are characteristic winged genera in the open country. Various papers on the
Apionine by Herr Hans Wagner, in which additional species from our region are
described, have been published during recent years. Unfortunately, very little
is known as yet of the actual food plants of any of the Central-American
Rhynchophora.
The fifteen coloured plates include figures of nearly 400 species.

24. CoteopTreRA. Vol. IV. part 4: by G. C. Champion: Curculionide (part).

The whole of this Volume is devoted to the Subfamily Curculionine, which are so.
numerous in the forest regions of Tropical America as to deter most Coleopterists
from venturing to describe them. Twenty-two groups are enumerated, the Sitonina
to the Cryptorrhynchina inclusive, numbering in all 1365 species, 1146 of which
are treated as new. Some of the genera include a very large number of species:
Conotrachelus (nearly 200), Anthonomus (over 100), Hilipus, Otidocephalus, Cryptor-
rhynchus, Kubulus, &c., so that it seems an almost hopeless task to prepare a complete
list of these insects. Dr. Sharp and Mr. Champion have been the first to describe
the whole of the Curculionide of a tropical country, and, though the genera of the
Cryptorrhynchina still remain in inextricable confusion, the present contribution will
doubtless be of considerable assistance to future workers. Some of the Groups,
Anchonina, Cholina, &c., are purely Neotropical ; others, Pissodina, Sitonina, Hyperina,
Balaninina, Cleonina (Lirus excepted), belong to more temperate regions, but extend
southward to within our limits.

The thirty-five plates include figures of nearly 1000 species: nineteen (x. and
XVill.—-xxxv. inclusive) are coloured, one (xii.) partly coloured, and the rest
uncoloured.

25. CoLEorTeRA. Vol. IV. part 5: by G. C. Champion: Curculionide (continued).

This Volume deals with four more Groups of the Subfamily Curculionine—the
Zygopina, Tachygonina, Ceuthorrhynchina, and Barina, the vast complex mass known
as Barina being represented by eleven Sections, all but three of which are purely
tropical. The Zygopina also are almost entirely tropical, a few forms only occurring
north of the Mexican frontier and they are wholly wanting in the European fauna..

70 CONTENTS OF EACH VOLUME.

The Ceuthorrhynchina, on the other hand, are numerous in temperate regions and
but poorly represented in the tropics. Of the 908 species enumerated for the four
groups mentioned, 620 belong to Barina; 717 are described as new, with 81 new
genera.

The remaining groups of the Curculionine—the Acamptina, Trypetina, and
Cossonina—are described in Vol. LV. part 7. The twenty-three plates belonging to
Part 5, illustrating 720 species, are wholly or partly coloured. |

26. Connoprera. Vol. IV. part 6: Brenthide by D. Sharp; Scolytide by WwW. X,
H. Blandford ; Anthribide by K. Jordan.

Three families of the Rhynchophora are enumerated in this Volume: the Brenthide,
published in 1895,.by Dr. Sharp; the Scolytide, published in 1895-1905, by
Mr. Blandford; and the Anthribide, published in 1906-1907, by Dr. Jordan.
Part 6 was reserved for them in 1895, long before the Parts 4 and 5 were commenced,
The general sequence of arrangement adopted in this work has therefore been
interrupted by the interposition of these families in the present volume; the
remainder of the Curculionine belonging to Part 5 were concluded in Part 7. The
total number of species included in the three families is: Brenthide 140 (104 new),
Scolytidee 272 (181 new), and Anthribide 193 (148 new). The Brenthide are mostly
confined to the forests of the tropics, and are particularly numerous in Tropical
America, Madagascar, and the Indo-Malay region. As regards the Scolytide the
author, Mr. Blandford, remarks that the genera are either cosmopolitan or Neotropical,
some of the latter being represented by stragglers in North America; he also says that
the southward limit of many northern forms is doubtless determined by the distribution
of the Conifer, which do not reach south of Nicaragua. . Dr. Jordan, in his intro-
ductory remarks on the Anthribide (p. 299), writes as follows :—‘‘ The present
treatise offers an illustration of the great increase in the number of known species
when a tropical district has been more exhaustively examined. Hardly three dozen
forms were known from Central America, while the material now before me comprises
close on 200 species.” ‘These insects are rarely found in large numbers, owing to their
ereat resemblance to the bark of trees on which they occur. The author divides the
Anthribide into two ‘subfamilies, the Pleurocerine and the Anocerine, and gives a
key to the whole of the genera (pp. 300-302). In the text the type of each genus is
indicated, and the species of the larger genera are tabulated.

Fourteen plates are issued with this volume: three for the Brenthide, six for
the Scolytide, and five for the Anthribide, the last mentioned being coloured.

27. Cotroptera. Vol. 1V. part 7: by G. C. Champion: Curculionide (concluded).

This Volume finishes the Rhynchophora, the rest of the Curculionide belonging to
the Subfamily Curculionine—the Groups Acamptina, Trypetina, and Cossonina—and

ZOOLOGY. 71

the whole of the subfamily Calandrine being here dealt with. An Appendix to the
Curculionide is added, on pp. 178-212, enumerating a few additional forms and
making some corrections to the synonymy. ‘The Cossonina and Calandrinz are each
represented in Central America by a large number of species, some of the Calandrids
attacking palms, cacti, sugar-cane, maize, Musacew, &c. The Mexican Cossonids (the
genus Cossonus excepted) had been previously studied by Wollaston and the Calandrids
by Chevrolat, nevertheless, with more abundant material, many new forms were found
in our collection. Altogether 344 species are enumerated, 231 of which are described
as new. ‘The nine plates are coloured or partly coloured.

28. CotropTerA. Vol. V.: Longicornia by H. W. Bates; Bruchides by D. Sharp.

The greater part of this Volume (pp. 1-436), published in 1879-1885, is devoted to
the enumeration of the Longicornia by Mr. Bates, the Bruchides, by Dr. Sharp,
published in 1885, occupying pp. 437-504. Altogether the two tribes number 1445
species: Longicornia 1273 (648 new), Bruchides 150 (117 new). Mr. Bates,
in his Introduction to the Longicornia, published in 1886, remarks as follows:
“Compared with the Tribe Geodephaga, it is beyond doubt far more numerously
represented in tropical than in extra-tropical lands, and its species and genera are
naturally multiplied in the highest degree in tropical forests, where woody vegetable
growths, to which the Longicornia are almost exclusively attached in their larval states,
are most numerous and varied. Although their beauty of form and colour has led to
their having been industriously collected, it is evident, from the number of new
species continually arriving from countries supposed to be fairly well explored, that we
are as yet far from possessing even an approximately complete knowledge of the whole
product of Nature in this department. ‘his is partly due to the recondite and, to a
great extent, nocturnal habits of a vast proportion of the species, and the difficulty of
the search for them in dense primeval forests where few clearings offer the necessary
openings.” The author thinks that the main conclusions arrived at after a similar
examination of the Geodephaga are confirmed, viz. (1) that the Central-American
fauna is essentially Neotropical ; (2) that the northern portion of the region (Mexico
and Guatemala) is not an extension southward of the Nearctic Province, but (3) that
it is a remarkably distinct subprovince of the Neotropical fauna, Dr. Sharp,
in his remarks on the Bruchides, numbering 150 in all, says (p. 437) that our
knowledge of these insects is not sufficiently advanced to enable any trustworthy
generalizations to be made in reference to the species found in Central America ;
and, as the northern parts of Mexico had been inadequately explored, it was not
possible to say what relationship existed between the North American species and
those of the regions southward.

73 CONTENTS OF EACH VOLUME.

Of the twenty-six coloured plates issued in the present volume, one only (pl. xxvi.)
is devoted to the Bruchides. .

Two papers on the Longicornia, subsequently received from our collectors after
Vol. V. was finished, have been published, entitled: ‘¢ Additions to the Longicornia of
Mexico and Central America”: one by Mr. Bates (Trans. Ent. Soc. Lond. 1892,
pp. 143-183, pls. v.—vii.); the other by Mr. Gahan (loc. cit. pp. 255-274, pl. xil.).
These additions bring the total number of species of Central American Longicornia
to 1372.

29, 30, Cornoptera. Vol. VI. part 1, and Supplement: by M. Jacoby: Phytophaga
(part).

The enumeration of the Phytophagous Coleoptera was commenced in 1880, before
the Sallé collection had been acquired by the Editors and shortly atter Mr. Champion
had started on his journey to Central America. ‘The coilections made by him, and
by Hage in various parts of Mexico, increased our material so largely that a Supple-
mentary Volume was added. Hence, two bulky Volumes, together including over 1000
pages of text and forty-three plates, were required for the account of the Families
Sagride, Donaciide, Crioceride, Megalopodide, Clythride, Cryptocephalide, Chlamy-
didz, Lamprosomide, Eumolpide, Chrysomelide, and Galerucide. The Hispide
and Cassidide are described in Vol. VI. part 2. In the general Introduction to the
two Volumes (Part 1 and the Supplement), published in 1892, the relative number
of species for each family is shown ina Table, and the number of species of the
larger genera (Diabrotica alone possessing 178, 116 of which are described as new,
and Lema 129, with 73 new) is also noted. At the end of this Introduction, a
systematic list of the whole of the species figured is given on pp. ix-xix. The Phyto-
phaga are perhaps more in evidence, except in the denser forest districts, than any of
the other families of Coleoptera in Tropical America, many of the species occurring
in great abundance on the herbage in open places at the commencement of the rainy
season. Some of them, like Ovina in Europe, are extremely variable in colour, so that,
as the author observes, nothing can be done by the systematic worker but to treat as
distinct such forms as in his opinion differ sufficiently from their allies. Of the 2166
species enumerated (in 1892) only about 90 are known from north of the Mexican
boundary and about 150 from south of Panama: Central America is thus shown to
have an exceedingly rich and peculiar Phytophagous fauna, of which the affinities are
much greater with South than with North America. Since the publication of these two
volumes, very little has been added to the Central American list beyond a few forms
described by Mr. Bowditch; Mr. Gahan’s papers on TRabrotica dealt with South
American species only. |

About 1000 species are figured on the forty-three coloured plates.

Pm ©

ZOOLOGY. i3

31. Corzoptera. Vol. VI. part 2: by J. S. Baly and G. C, Champion: Phytophaga
(part).

This Volume gives an account of the Hispide by J. 8. Baly, published in 1885-1886,
with a short Appendix, in 1894, by G. C. Champion; and of the Cassidide by
G. C. Champion, published in 1893-1594. For the two families 453 species are
enumerated—Hispide 226 (169 new), Cassidide 227 (59 new). The Central-
American representatives of these groups are essentially N eotropical, America north
of Mexico possessing a very limited number of species. Many of the characteristic
South-American genera of Cassidide, however, do not reach so far north as Panama,
or, if present, are represented by very few species. Mexico is particularly rich in
peculiar forms belonging to the genera Che/ymorpha, Physonota, Coptocycla, &c.
‘The Hispidee attack Musacee, Bambusacee, &c., and abound throughout the warmer
parts of Tropical America.

The thirteen coloured plates inclnde figures of nearly 300 species.

82. CoLeoptera. Vol. VIL: by H. S. Gorham: Erotylide, Endomychide, and
Coccinellidee. .

The three families enumerated in this Volume are placed by recent writers in the
Clavicorn-series, which are dealt with in Vol. II. part | of the present work. The
total number of species is as follows:—Erotylide 282 (154 new) ; Endomychidee
81 (39 new); and Coccinellide 239 (108 new). The Erotylide, including the
Languriides, were published in 1887-1889, the Endomychice in 1889-1891, the
Coccinellide in 1891-1898, and a short Supplement to all three families, and the
Introduction, in 1898-1899. The Author, speaking of the Erotylide, exclusive of
the Languriides, says that the members of this highly developed family of fungivorous
beetles are largely endemic. The Endomychide are somewhat poorly represented in
Central America; the Coccinellide, or ‘lady-birds,’ on the other hand, are very
abundant in species and individuals, the phytophagous Epilachne in particular. The
small Aphid- and Coccid-devouring Coccinellids have of recent years become of

economic importance.
The thirteen coloured plates include figures of nearly 300 species.

This is the last of the eighteen volumes devoted to the Coleoptera; the total
number of species enumerated is 18,029. Several genera of doubtful position, left
undetermined by the various authors, have recently been described and figured, and
the species of the genus Hapalips revised, by Mr. Champion (Trans. Ent. Soc. Lond.

1913, pp. 58-169, pls. iii, iv.).

83 Hymenoprera. Vol. 1.: By P. Cameron: Sessiliventria and Petioliventria.

The fifteen families of Phytophagous or parasitic Hymenoptera worked out in this
Volume number in all 1109 species, of which 596 are described as new. The

BIOL. CENTR.-AMER., Introd. Vol., January 1915. L

74 CONTENTS OF EACH VOLUME.

Sessiliventria include the Tenthredinide, Cephide, Siricide, and Orysside, and the
Petioliventria the Cynipide, Figitide, Chalcidide, Ichneumonide, Braconide,
Stephanide, Evaniide, Trigonalide, Pelecinide, Proctotrupide, and Chrysidide.
When Mr. Cameron’s work on these insects was published, in 1883-1888, scarcely any
Mexican material was available for study, and the numerous Ichneumonide described
from that country in 1873 by Cresson could not therefore be identified. Mr. H. H.
Smith subsequently made very extensive collections in Mexico of all the above-
mentioned families, too late, unfortunately, to be included. Mr. Cameron’s contri-
bution, therefore, is very incomplete as regards the total number of species inhabiting
the region, although it adds considerably to our knowledge of these Hymenoptera.
The numbers of species for the different families are as follows: Tenthredinide 152,
Cephide 1, Siricide 3, Orysside 2, Cynipide 15, Figitide 9, Chalcidide 121,
Ichneumonide 533, Braconide 176, Stephanide 5, Evaniide 23, Trigonalide 9,
Pelecinide 2, Proctotrupide 43, and Chrysidide 19.
The twenty coloured plates include figures of 451 species.

34, Hymenoptera. Vol. II.: by P. Cameron: Fossores.

In this Volume the species belonging to the Section Fossores (the sand- and wood-
wasps), under which the author includes the Family Mutillide, of the Hymenoptera
Aculeata, are enumerated; the other Sections are known as Heterogyna (ants), Diplo-
pterygia (Eumenide and Vespide), and Anthophila (bees). ‘The Fossores, including
Mutillide, number 711 species, of which 391 are treated as new, the total number
being: Sphegide 67, Ampulicide 2, Larride 52, Nyssonide 382, Bembecide 26,
Philanthide 54, Mimeside 8, Pemphredonide 3, Crabronide 24, Pompilide 163,
Scoliidee 59, Mutillidee 221. The pages 1-400 were published between 1888-1896,
the rest in 1899 and 1900. Mr. H. H. Smith’s Mexican collections fortunately
arrived in time to be included, so that the enumeration of the species in Vol. II. is
more complete than in Vol. I. of the Hymenoptera. In the Introduction (published

in 1900) there is a classified list of the 312 species figured on the fourteen coloured
plates.

35. Hymenoptera. Vol. III.: by Prof. A. Forel: Heterogyna.

Volume III. of the Hymenoptera, published in 1899-1900, contains an account, in
French, of the Heterogyna or Ants, numbering 382 species, 66 of which are described
as new. From the author’s remarks on page 1, it would seem that a large number
of the Central-American forms are common to South America, and that the Formicid-

fauna is mainly Neotropical. A few closely related S.-American species are described

in foot-notes. The four uncoloured plates include figures of 57 species ; plate II. shows
the nests of various ants among leaves, or in the spines of Erythrina, Acacia, &. A
considerable number of species have been added to the fauna by Prof. Forel and other
writers during the past ten years, showing that new forms still await the collector.

ZOOLOGY. 75

[The rest of the Aculeata, including the social-wasps, bees, etc., remain unworkee,
no contributor having been willing to undertake the large amount of material obtained
in these groups. The late Lieutenant-Colonel Bingham commenced the study of the
Diploptera shortly before his death, but nothing was published. ]

36-38. LeprpopreRA Ruopanocers. Vols. I., IL. (text), III. (plates): by F. D. Godman
and QO, Salvin; with (in Vol. II.) a Note on the Group Eumeidi by
S. H. Scudder.

Vol. I. of the Butterflies, published in 1879-1886, the Introduction excepted, gives
an account of the Nymphalide—including the subfamilies Danaine, Satyrine,
Morphine, Brassoline, Acreine, Heliconiine, and Nymphaline,—Libytheide, and
Erycinide. Vol. IL. includes the enumeration of the Lyceenide (published in 1887),
Papilionide, including the subiamilies Pierine and Papilionine (1889-1893), and
Hesperiide (Hesperiine, 1893-1899, Pamphiline, by Godman alone, 1900-1901); a
Supplement to the whole subject by Godman, pp. 638-741 (1901); and a Note on the
Group Eumeidi, pp. 110-112, published in 1887, by Scudder. The Introduction to
Vol. I. (pp. v-xlvi), by Godman, issued on the conclusion of Vol. II., in 1901, contains a
description of the physical conformation of each of the countries belonging to the Central-
American region; an explanation of the classification adopted ; remarks on the leading
forms belonging to each of the six families recognized (Nymphalide, Libytheide,
Erycinidw, Lycenide, Papilionide, and Hesperiide) ; a Table showing the geographical
distribution of the genera (pp. xxli-xxvii) ; particulars as to the sources from whence
the material was obtained, with the names of the collectors and some remarks on the
places visited by them; and a systematic list of the whole of the species figured (1206)
in the two Volumes (pp. xxxi-xlv), the plates numbering 112 in all. Altogether, 1805
species (Nymphalide 588, Libytheide 1, Erycinide: 240, Lycenide 234, Papilionide 186,
and Hesperiide 556) are enumerated, 376 of which are described as new, and 18 others
from South America are added in foot-notes. In the Hesperiide the genitalia of most
of the species are figured, many closely allied forms being definitely separable by these
male-structures, and some of the species of the genus Papilio are similarly illustrated.

The study of these insects is said to prove conclusively (1) that the fauna is mainly
a northern extension of that of Tropical South America, extending on the Pacific side
to Mazatian in Mexico and on the Atlantic side to a little beyond Ciudad Victoria
in Tamaulipas, with many peculiarly modified forms in the region; (2) that
there are a considerable number of Nearctic genera and species coming down the
central plateau a certain distance into Mexico, and some even to Guatemala ; (3) that
there are no strictly alpine forms; (4) that the fauna of the Atlantic slope to perhaps
as far south as Costa Rica is incomparably richer than the Pacific; (5) that some of
the purely tropical genera do not reach north of Nicaragua, Costa Rica, or Panama.

The publication of the concluding portion of the Rhopalocera, as in the case of the
L2

76° CONTENTS OF EACH VOLUME.

Aves, was greatly delayed by the ill health and death of Salvin, in 1898; but he lived
long enough to assist in completing the enumeration of the species of nearly all the
families, with the exception of those of the subfamily Pamphiline of the Hesperiide,
i. é. to p. 460 of Vol. LI.

Amongst the numerous papers on Neotropical Butterflies that have been published
since the conclusion of Vol. II. (in 1901), one by Mr. H. H. Druce (P. Z. 5. 1907,
pp. 566-632, pls. xxxi—xxxvi.) contains a revision of certain Central-American species
of the genus Thecla.

39-41. Leprpoprera Hurerocers. Vols. L., IL. (:ext), I11. (plates): by Herbert Druce :
Sphingidee—Pyralide.

Vol. I. of this subject, published in 1881-1891, the Introduction excepted, contains
an account of the following 22 families of Moths :—Sphingide, Castniide, /Egeriide,
Agaristide, Zygenide, Arctiide, Chalcosiide, Lithosiide, Melameride, Dioptide,
Liparide, Saturniide, Lasiocampide, Limacodide, Bombycide, Drepanulide, Psychide,
Cosside, Hepialide, Notodontide, Noctuide, and Deltoide. Vol. II., published in
1891-1896, includes the Euschemide, Uraniide, Geometride, Siculide, and Pyralide ;
and a very extensive Supplement, pp 298-569, issued in 1896-1899, embraces all
the families contained in both Volumes, another family, the Arbelide (pp. 449, 450),
being added. The author, in his Introduction to Vol. I., published in 1900, states
that in 1880 less than 400 species of Heterocera had been recorded from Central
America against the 3639 enumerated in these two Volumes. The Table given
on page ix shows the number of species belonging to each of the 28 Families and
of those peculiar to the region. He says that it is almost impossible to give any
generalizations as to the composition of the Central American Heterocera, because so
little is known of the species inhabiting the adjacent parts of South America; but it
may safely be said that their affinities are almost entirely South American, and the
few northern forms that do occur mostly belong to widely distributed genera.

The 101 coloured plates include figures of 1926 species, a complete list of which is
given in the Introduction to Vol. I., pp. xi-xxxi.

Since the conclusion of Mr. Druce’s work, in 1900, a large number of species have
been added, especially from Mexico and Costa Rica, mainly by Mr. W. Schaus, who
has specially visited Costa Rica, &c., in search of Lepidoptera. Sir George Hampson,
too, in his revision of the Noctuide, Pyralidz, &c., has also contributed largely to our
knowledge of the Tropical American Heterocera, the critical study of which had
scarcely been commenced when Vol. I. was undertaken.

42, LepipopTeRA Hererocera. Vol. IV. (text and plates): by Lord Walsingham :
Tineina, Pterophorina, and Orneodina.

This Volume, commenced in 1909, contains the enumeration of the ‘Tineina,

ZOOLOGY. TT

Pterophorina, and Orneodina, and a few supplementary genera and species omitted in
the other volumes dealing with the Heterocera. The Tineina are grouped under 22.
Families—Lavernide, Gelechiade, GEcophoride, Ethmiade, Blastobaside, Stenomide,
[ Aigeriade], [Cosside], Arrhenophanide, Sparganothide, Olethreutide, Tortricide,
Phaloniade, Carposinide, Hemerophilide, Coleophoride, Heliodinide, Hypono-
meutide, Phyllorycteride, Tineidz, Acrolophide, and Nemophoride. ‘Two of them,
the Aigeriade and the Cosside, were previously dealt with in detail by Mr. Druce
in Vol. I., their true affinities with the Tineina not having been recognized at
that time. Lord Walsingham, in his Introduction, alludes to the great assistance
he has received from Mr. J. H. Durrant in the preparation of this Volume, which
is illustrated by ten coloured plates, representing 350 species of Tineina.

43, Diprera. Vol. I.: by Baron C. R. Osten Sacken, the Supplement by S. W. Wil-
liston, J. M. Aldrich, W. M. Wheeler, and A. L. Melander: Cecido-
myiicdee—Em pide.

Twenty-four Families of Diptera are enumerated in this Volume—Cecidomyiide,
Mycetophilide, Bibionide, Simuliide, Blepharoceride, Culicide, Chironomide,
Psychodide, Tipulide, Rhyphide, Stratiomyide, Tabanide, Chiromyzide, Leptide,
Xylophagide, Acanthomeride, Mydaide, Nemestrinide, Bombyliide, ‘Therevide,
Cyrtide, Asilide, Dolichopodide, and Empide. Osten Sacken’s work was published
in 1886-1887. ‘The Supplement by the American writers Williston, Aldrich, Wheeler,
and Melander was issued in 1900-1901, and the editorial Introduction in 1901, on
the conclusion of the whole subject. The material examined by Osten Sacken was
very meagre, and though more specimens from Mexico were available for the
Supplement, yet it was insufficient to give more than a general idea of the Dipterous
fauna of Central America. The present Volume, therefore, is merely a contribution
towards the study of these insects, rather than a complete list of the forms actually
inhabiting the region. Altogether, 977 species are enumerated, 330 of which are
described as new. ‘The six coloured plates illustrate 120 species.

During recent years the Mosquito fauna of Panama has been specially studied in
the canal-zone by Mr. A. Busck, and his report has been published [Smithsonian
Contributions, vol. lii. pp. 49-77 (May Ist, 1908)]. Many uew species of Culicidee
from Panama and other places in Central America have been described by Theobald,
Dyar, and Knab.

44, Diprera. Vol. Il.: by F. M. van der Wulp: Qiéstride, Muscide, and Hippo-
boscide.

This Volume contains the enumeration of the numerous groups of Diptera known
collectively under the name of Muscide, for which abundant Mexican material collected
by Mr. H. H. Smith was available, but the work was only about half finished when the

78 CONTENTS OF EACH VOLUME.

author died in 1899. No other contributor could be found to undertake the groups
Agromyzine, Borborine, Chloropsine, Drosophiline, Ephydrine, Geomyzine, and
Sapromyzine, of the Muscide Acalyptere, and the Phoride, and the volume was
closed as it stood. Pages 429-489, containing the Hippoboscide and the Supple-
ment to the other families, were published after v. der Wulp’s death, although
the new species of Muscide Calyptre had already been described by him in the
‘Tijdschrift voor Entomologie’ for 1892. Altogether, 1095 species were named by the
author, of which 585 were treated as new, the publication of his work extending from
1888-1900, but the Titlepage and Introduction were not issued till 1903.

The thirteen coloured plates include figures of 287 species. Some of the Trypetine
are figured on the plates, others (11) in the text.

45, Dirrera. Vol. III.: by S. W. Williston: Syrphide, Conopide, Pipunculide, and
Platypezide.

Prof, Williston’s contribution to this Volume, pp. 1-89, was published in 189i-
1892, and the list of subsequently described species, and the general index to the
whole of the three volumes, pp. 93-127, in 1903. The material again was scanty in
comparison with the large number of species which must exist in the region, but
among the Syrphids the genera Baccha, Volucella, and Eristalis were particularly
well represented. The total number of species enumerated is 320, of which 67
are new. The two coloured plates include figures of 29 species.

46. Ruyncnota Hereroprersa. Vol. I.: by W. L. Distant: Pentatomide, Coreide,
Lygeide, Pyrrhocoride, and Capside.

Mr. Distant’s enumeration of the above mentioned families of Rhynchota Hetero-
ptera (pp. 1-303) was issued in 1880-1884, the Supplement to the same (pp. 304-351)
in 1889-1893, and the Appendix (pp. 452-462) in 1893, the delay in publishing
the Supplement being due to his long absence in South Africa, Altogether,
1108 species are recognized, more than half of which are described as new. In the
Introduction, published in 1893, the number of genera belonging to each family is
given and the geographical distribution shown in short Tables. Amongst the
Lygwide nearly half the genera are stated to be endemic. The Pentatomide include
377 species and are, therefore, very well represented in Central America, though
many of the genera are common to the Neotropical region. The Capside, too, with
313 species, are very numerous, but the tropical representatives of this family are
but little known as yet, and no comparison with the fauna of the adjacent portion
of South America can be made. The thirty-nine coloured plates include figures of
upwards of 900 species, a complete list of them being given in the Introduction,
pp. Xi-Xx.

ZOOLOGY. 79

47. Ruyncnota Hetrrorrera. Vol. II.: by G. C. Champion: Tingitids-Corixide.

Mr. Champion here deals with nineteen families of Rhynchota Heteroptera (making
twenty-four in all), twelve belonging to the Gymnocerata and seven to the Crypto-
cerata. The number of species enumerated is 592, nearly half of which are
described as new. Fifteen additions to Vol. I., described or recorded by various
authors in other works, between 1893 and 1901, are noted on p. 384, the total
number of Heteroptera enumerated in the two Volumes being 1715. The Tingitide,
Aradide, Reduviide, and Anthocoride are abundantly represented in Central America,
the respective numbers for each being 78 (66 new), 78 (40 new), 204 (78 new),
and 54 (32 new). As with the five families belonging to Vol. I., no comparison is
made with the fauna of the adjacent regions southward for want of sufficient data.
Most of the genera belonging to the aquatic Cryptocerata are, in common with other
water-insects, more or less widely distributed. The twenty-two plates, six of which
are coloured, include figures of 513 species, a complete list of which is given in the
Introduction, pp. xi-xvi. The Index for the two volumes is included in Vol. IL,
pp. 385-416.

48. Ruyycnota Homoptera. Vol. 1.: by W. L. Distant and W. W. Fowler: Cicadide—
Delphacide.

Mr. Distant’s first contribution to this Volume, on the Cicadide and Fulgoride,
pp. 1-41, was published in 1881-1887, the Supplement to the same, pp. 42, 43, in
1900, and the Appendix, pp. 140-146, in 1905, his work having been interrupted
during long absences from England. The rest of the volume, dealing with three
additional species of Dictyopharine, the Flatide, Derbide, Cixiidw, Achilide, Isside,
and Delphacide, by the Rev. W. W. Fowler, pp. 44-139, 146, 147, appeared in
1900-1905. Apart from the Cicadide and Fulgoride, very little appears to be known
about the South American members of the above mentioned families, and nearly all the
Central American representatives are described as new by Mr. Fowler. Altogether 323
species are enumerated, 208 of which are treated as new, with twenty-five new genera.
The thirteen coloured plates include figures of 267 species, a complete list of which
is given in the Introduction, pp. vii-ix. Four species of Cicadide are also figured
in the text, on pp. 140-143. A few Central American Cicadide have since been
described by Mr. Distant in other publications.

49, Ruyncnota Homoprera. Vol. II. Part 1, by W. W. Fowler: Membracide-
Gyponide. Part 2, by T. D. A. Cockerell: Aleurodide and Coccide.

Part 1 of this Volume contains the enumeration of four more families of Homoptera
—the Membracide, Cercopide, Tettigoniide, and Gyponide, 659 species in all
(exclusive of the 29 noticed in the Supplement, some of which are doubtful), 385 of
of these being described as new, with 42 new genera. Pages 1-316 were published

80 CONTENTS. OF EACH VOLUME.

in 1894-1903, and 317-322 in 1909, the conclusion of the Volume having been
delayed for several years by the American contributer who had volunteered to
determine the Jasside, Bythoscopide, &c. The MSS. and material not being forth-
coming, the volume was closed as it stood. ‘The twenty-one coloured plates, ten
of which are devoted to the extraordinary insects known as Membracide, include
figures of 494 species, a list of which is given in the Introduction, pp. vii-xi. The
Index for Vols. L. and II. is also issued in this volume, pp. 823-339. The Supplement,
pp. 818-322, contains notes on the 29 species added by Buckton and others during
the progress of the work.

Part 2 of this Volume, by Prof. T. D. A. Cockerell, issued in 1899 (the separate
Index, pp. 35-37, was not published till 1909, when Volume II. was closed}, gives
a list of the 161 Aleurodide and Coccide from Mexico and Central America known
to the author at that date. Eleven species and two genera are described as new,
and nive forms are figured in the text. With these insects, the total number of
Homoptera recorded from Central America is 1143.

[The few specimens of Thysanoptera obtained have been examined, and some of
the species described, by Mr. R. S. Bagnall (Journ. Linn. Soc., Zool. xxx. pp. 869-
387, pls. ii., iii, 1910). Mr. D. L. Crawford has also recently written various papers
on ‘Thysanoptera of Mexico and the South’ (Pomona Jonrn. Ent. i., ii. 1909-1910). |

50. Nevroprera: Ephemeride by A. E. Eaton; Odonata by P. P. Calvert.

The Rev. A. E. Eaton’s account of the Ephemeride, based upon very few specimens,
was published in 1892; that of the Odonata by Prof. P. P. Calvert, with abundant
material, in 1901-1908 *. Altogether 308 species are enumerated: Ephemeride 32
and Odonata 276. Prof. Calvert’s Introduction, pp. v-xxx (published in 1908),
mainly consists of two Tables: (A) ‘ Alphabetical List, by countries, of the localities
in which the Odonata were collected,’ and (B) ‘‘ Systematic list and distribution of
the species and varieties,’ 293 in all; anda coloured map to show the actual distri-
bution of mean annual temperatures. ‘Table A gives the name of the Department,
Territory, or State to which each locality belongs, the elevation of these places in
metres and feet, the names of the collectors, the dates of capture, general remarks, &c.
Table B shows the geographical distribution and the temperature zones in which
the species occurs in each country, this being further illustrated by the map. The
composition and ecological relations of the Odonate fanua of Mexico and Central
America have been discussed by Mr. Calvert elsewhere [ Proc. Acad. Nat. Sci. Phil.
(1908) 1909, pp. 460-491, pl. xxvi.]. He has since visited Costa Rica, May Ist,
1909-May 9th, 1910, to collect data on (1) the seasonal distribution of the Odonata,

* The Odonata were undertaken successively by Mr. McLachlan and Prof. Karsch, before Prof. Calvert
procceded with the work.

ZOOLOGY. $1

(2) their larval forms, and (3) their habits. An account of his journey was published
in the ‘ Entomological News’ for July 1910, pp. 334-337, and this has been followed
by other papers on the results of his researches, the most important being the
discovery of Odonate larve found living in the water which collects between the bases
of the leaves of Bromeliads growing upon trees, and of the rearing of the imagines of
Mecistogaster modestus from these larve.

The ten plates belonging to this Volume, of which three are coloured, include
figures of 178 species, all but nine appertaining to the Odonata.

[No contributor has been found for the remaining families belonging to the
Neuropterous-series in the wide sense, viz. Trichoptera, Neuroptera Planipennia,
Mallophaga, Psocidze, Termitide, Embiide, and Perlide. |

51. Ortuoprera. Vol. 1.: Forficulide by Count de Bormans; Blattide, Mantide,
Gryllide, and Locustide by Dr. Henri de Saussure, assisted by Dr. Leo
Zehntner and A. Pictet.

Five families of Orthoptera are dealt with in this Volume: the Forficulide by
de Bormans (published in 1893), and the Blattide, Mantide, Gryllide, and Locustide
by de Saussure (published in 1895-1899). The Introduction, including the systematic
list of species figured, was issued in 1900, when Vol. IL. of this subject was com-
menced. De Saussure, in his enumeration, also considered it necessary to describe,
notice, or figure various allied South-American or Antillean forms. | Under the
heading to each family, tribe, &c., he deals with the general classification of the
Neotropical fauna, and gives synopses of the genera, and under the genera them
selves dichotomous keys to the species. The total number of species recognized as
belonging to Central America is 633, of which 224 are new: Forficulide 41
(8 new), Blattide 156 (60 new), Mantidze 61 (33 new), Gryllide 102 (43 new),
Locustide 273 (80 new). The twenty-two plates, of which, iii., vi., vil., and viii.,
are coloured, include figures of 310 species.

52. OrtHoprerRa. Vol. II.: Acridiide by Prof. Lawrence Bruner (the subfamily
Tettigine by A. P. Morse); Phasmidee by R. Shelford.

This Volume includes the enumeration of the two remaining families of the
Orthoptera. The Phasmide should have been placed between the Blattide and
Mantide, in Vol. I., but the account of them had to be deferred until the appearance
of Brunner vy. Wattenwyl’s and J. Redtenbacher’s Monograph, the concluding portion
of which was not published till 1908. Prof. L. Bruner’s work on the Acridiide, in
which Mr. A. P. Morse assisted him with the subfamily Tettigine, was issued in
1900-1908, the Tettiginee having appeared in 1900-1901. ‘The list of the Phasmide,
published in 1908, was compiled from the above mentioned Monograph, the authors

BIOL. CENTR.-AMER., Introd. Vol., “une 1915. M

82 CONTENTS OF EACH VOLUME.

of which included the material obtained by the Editors of the present work. Asin
Vol. I., the species, excepting the Tettigine, likely to occur in Mexico or Central
America are included in the enumeration. Altogether 663 species are regarded
as belonging to the fauna, although in many cases, owing to lack of material, there
is at present no actual record from within the limits of Central America. The 663
species are apportioned thus: Acridiide, including the 26 Tettigine, 522 (138 new) ;
Phasmide 141. The Index to the two volumes occupies pp. 379-412 in Vol. II.
‘The eight uncoloured plates (four for each family) include figures of 120 species, some
of which are Nearctic or Neotropical forms closely related to those known to inhabit
Central America.

BOTANY.

53. Vol. I.: by W. B. Hemsley, with a Commentary on the Introduction and Appendix
by Sir J. D. Hooker.

This Volume contains the enumeration of the Polypetale, Ranunculacee-
Ayaliacese, which was issued in 1879-1881; the Preface, Introduction, and ‘Com-
mentary ’ appeared in October 1888, on the completion of the entire subject. When
this took place, the temporary titlepage and the Index to Vol. I. (pp. 577-619),
issued in October 1881, were both replaced in October 1888, the former by a titlepage
to include notice of the ‘Commentary’ by Sir J. D. Hooker, and the latter
by a general index for the whole of the four Volumes of text, this appearing in
Vol. IV. pp. 833-498, 1887-1888. In the Introduction, pp. ix—lxi (issued in 1888),
Mr. Hemsley deals with geographical distribution under the following headings:
(1) Statistics of the Phanerogamic Flora of the world; (2) Statistical comparison of
the Floras of large and widely separated areas; (3) Generic and specific composition
of the Floras of different areas; (4) The distribution of some of the largest Natural
Orders; (5) The Primary Botanical regions of the World considered in their relations
to the Zoological regions; (6) Comparison of the Zoological with the Botanical
regions; (7) Outlying Australian types of vegetation; (8) Botanical division of the
earth into primary regions. Sir J. D. Hooker in his ‘Commentary on Mr. Hemsley’s
Introduction and Appendix to the Botanical part of the Biologia Centrali-Americana’
(pp. lxii-xlviii) also gives his own views on the Botanical kingdoms, under these
headings: (i.) The North Temperate Kingdom of the Old World; (ii.) The Tropical
Kingdoms of the Old and New World; (iii.) The three Southern temperate regions
(Extratropical America, Africa, and Australia).

54. Vol. II.: by W. B. Hemsley.

The Gamopetale, Caprifoliacee—Plantaginee, are dealt with in this Volume. The
temporary titlepage and the Index to Vol. II., pp. 577-621, both issued in June

BOTANY. 83

1882, were cancelled in 1888, the ‘“‘ Contents” being added to the title and the Index
replaced (as in Vol. I.) by one for the complete series of four volumes, this being
inserted at the end of Vol. IV.

55. Vol. III.: by W. B. Hemsley ; the Cycadacee by W. T. Thistelton-Dyer.

The Incomplete, Monocotyledones, and Cryptogamic Vasculares, Nyctagina—
Rhizocarpe, are enumerated in Vol. III. The Cycadacee, pp. 190-195, were studied
by Mr., now Sir W. T. Thistelton-Dyer, and his portion of the subject was issued
in 1883.

56. Vol. 1V.: by W. B. Hemsley.

This Volume, the last of the series, exclusive of that occupied by the Plates.
includes: (1) The Supplement to Vols. I.-III.; (2) The enumeration of a small
collection of plants made in Cozumel Island in 1885; (3) A list of plants from
Holbox, Mugeres, Cozumel, and Ruatan Islands made in 1886; (4) Additions to the
list of Costa Rican ferns, bringing the number up to 134; (5) Appendix. The
Appendix deals with a variety of subjects: (i.) Preliminary remarks; (ii.) A sketch of
the history of the botanical exploration of Mexico and Central America ; (iii.) Outlines
of the geography and the prominent features of the Flora of Mexico and Central
America ; (iv.) Summary and analysis of the Flora; (v.) Relationships with the Floras
of other regions; (vi.) Further details of the distribution of some of the more
prominent natural Orders ; (vii.) A specimen of the mountain Flora of South Mexico
and Central America; (viii.) Altitudinal distribution of Orchids in South Mexico
and the dominating features of the general vegetation; (ix.) Recapitulation of the
dominant features of the Flora of Mexico and Central America, and remarks on its
probable derivation ; (x.) Bibliography. ‘The general index, as stated above, occupies
pp. 333-498. The total number of species enumerated in the four volumes is
11,626, of which 196 are indicated as new by the authors.

57. Vol. Vv. : by W. B. Hemsley.

The 111 Plates (including XLI.*), eighteen of which are coloured, together form
Vol, V. of this subject. A complete list of the 144 species figured is given on
pp. v-viii. Plate CX. consists of a Map of Mexico and Central America, showing the
then known northern limits (in 1887) of the Phanerogamic epiphytes, the Coco-palm,
the Marcgraviacee, the Vochysiacez, and the scuthern limit of Pines.

84 CONTENTS OF EACH VOLUME.

a ARCH AOLOGY.
58, 59. Vol. I. (text and plates): by A. P. Maudslay. |

The Volume of text contains the Preface and Introduction to the entire subject,
and a general account of the ruins at Copan. The latter are described under five
headings: (1) Principal Notices and descriptions of the Ruins; (2) Personal narrative ;
(3) General description of site; (4) Detailed description of principal structures ;
(5) Description of Stele and Altars. In addition to the 119 folio plates, there are
numerous illustrations in the text. The text of Vols. I-IV. is arranged for binding
in one Volume; the plates illustrating the subject form four separate volumes,
when bound.

58, 60. Vol. IL. (text and plates): by A. P. Maudslay.

The Ruins described in Vol. II. are those of Quirigua, Ixkun, Yaxché, Rabinal,
Chacujal, Utatlan and Iximché, Mixco (foundation-mounds), and Menché. These
places are all shown on the 98 folio plates, the first of which is a map of Guatemala
and the adjoining countries, showing the position of the ruins; and there are also
various illustrations in the text.

58, 61. Vol. III. (text and plates): by A. P. Maudslay.

The Ruins described in Vol. III. are those of Chichén Itza and Tikal. The 82 folio
plates, the first of which is a map of Yucatan and the country to the south of it, show
these places, and there are also three illustrations in the text.

58, 62. Vol. IV. (text and plates): by A. P. Maudslay.

The Ruins of Palenque are described in Vol. IV. under eight headings: (1) Personal
Narrative; (2) Principal Notices and Descriptions of the Ruins; (3) General de-
scription of the site; (4) Detailed description of the principal structures; (5) Pottery;
(6) Initial Series or date; (7) The two-headed dragon ; (8) The Water-plant. These
subjects are shown on the 98 folio plates.

63. Appendix (text only): The Archaic Maya Inscriptions: by J. T. Goodman.

Pages 1-149 of this Volume are occupied with Mr. Goodman’s account of these
inscriptions; and pp. 151-264 by the Annual, Chronological, and Perpetual Calendars

(Tables), and the Working Chart. The ‘Appendix’ forms the entire Part VIII. of
the Archeology, and is unaccompanied by Plates.

SUMMARY OF CONTENTS OF EACH VOLUME.

Total New
number of! species
species. | described.
1. Introductory Volume ..........
ZOOLoey.
2. Mammalia .................... 181
3. Aves. Vol. I. ..............
4, ow > Vol. TE. ...........8,. 1413 14
5. » Vol. Id. ..............
6. yo VOL TV. owe. eee eee .. ws
7. Reptilia and Batrachia.......... 695 58
S. Pisces 2c. cee eee eee 415 16
9. Terrestrial and Fluviatile Mollusca] 871 81
10. Arachnida Araneidea .......... 1181* 417
11. Aelia - and Opiliones. | { 422+
achnida Seorpiones, Pedipalpi,
ee and Solifugee ° tec eeees , . . . } 69 8
13. Arachnida Acaridea ............ 5d 40
14. Chilopodaand Diplopoda........ 255 106
15. Coleoptera. Vol. I. part 1 ....| 1086 384
16, » Vol. I. part2 ....| 1790 147
17. ” Vol. Il. part 1 ....} 1629 996
18. » Vol. II. part2 ....) 1100 494
19. 9% Vol. III. part 1 ....| 1358 805
20. i. Vol. III. part2 ....| 818 5B5
21. a Vol. IV. part 1 .... —
22, Vol. IV. part2 .... } 1776 | 1295
23. a Vol. IV. part3 ....) 616 516
24. % Vol. IV. part 4 1365 1146
25. $9 Vol. IV. part 5 908 717
26. ” Vol. IV. part 6 605 433
27. is Vol. IV. part 7 344 231
28. 55 Vol. V. ws... .e eee 1423¢ 765
29. 7 Vol. VI. partl .... ]
30. oa Vol. VI. part 1, 2166 1863
Supplement .. f
ol. ” Vol. VI. part 2.... 453 224
32. . Vol. VIL........... 602 301
33. Hymenoptera. Vol. I. ...... 1109 596
34. - Vol. ID ...... 711 391
35. - Vol. UI. ...... 382 66
36. Lepidoptera Rhopalocera. Vol. I
37, ; Vol. II. } 1805 376
38. "2 - Vol. III}... ..
39. :. Heterocera. Vol. LI.
oe 7 : aes | 8639 1312
41. = 4 Vol. IIL)... =
42. a " Vol. IV) 973$ 586

New Plates Speci Dates
ener includi pecies Pages of of
d & oribed, ( maps). figured. letterpress. issue.
10 vili & 149 1915
22 34 xx & 220 1879-1882
xliv & 512 | 1879-1904
1 iv & 598 1888-1904
iv & 510 1897-1904
.. 84 149 vil 1879-1904
3 76 138 xx & 326 1885-1902
1 28 78 | xxxiii & 2038 | 1906-1908
1 dt 333 | xxvili & 706 | 1890-1901
4 39 981 xv & 317 1889-1902
38 o4 1 x & 610 1897-1905
12 37 i& 71 1902
.. 21 43 xxi & 55 1886-1893
12 15 139 1 & 217 1895-1910
8 13 324 x & 316 1881-1884
103 19 443 xvi & 824 1882-1887
83 19 475 xii & 717 1887-1905
19 24 492 xu & 482 1886-1890
28 27 648 xvi & 690 1882-1897
28 13 330 xii & 372 1880-1886
99 { 23 568 | xxxiv & 572 | 1884-1893
- 21 500. x & 494 1889-189:3
45 15 397 vi & 354 1889-1911
69 35 997 vili & 750 1902-1906
81 23 720 vill & 514 1906-1909
32 14 324 iv & 396 1895-1907
17 9 275 vi & 221 1909-1910
57 26 515 xii & 526 1879-1886
xx & 625 1880-1892
43 43 | 1010 \ iv & 874 | 1888-1892
ae 13 291 x & 250 1885-1894
21 13 287 xii & 276 1887-1899
21 20 451 xii & 488 1883-1900
1 14 312 xi & 413 1888-1900
+. 4 57 1 & 170 1899-1900
119 f xlvi & 487 | 1879-1901
_ nl iv & 782 1887-1901
.. 112 1206 iv 1879-1901
4l xxxii & 490 | 1881-1900
_ - iv & 622 1891-1900
.. 101 1926 iv 1881-1900
o4+ 10 350 xii & 482 1909-1915

* Araneidea 1111, Opiliones 70.

+ Araneidea 364, Opiliones 58.
+ Longicornia 1273 (648 new), Bruchides 150 (117 new).
§ Exclusive of 52 species previously enumerated by Druce in Vols. I. and II.

86

SUMMARY OF CONTENTS OF EACH VOLUME.

Total New New Plates Speci » f Dates
: . . pecies ages 0 of
ranere: | ackenbed. | afseribed. |“'maps). "| Seared. | letterpress issue.
ZOOLOGY (cont.).
43. Diptera. Vol. I. ............ 977 330 11 6 120 viii & 378 1886-1901
44, a Vol. IL. wo... ee eee eee 1095 585 40 13 287 x & 490 1888-1903
45. » Vol ld. .........e.. 320 67 1 2 29 | vili& 127 | 1891-1903
46. Rhynchota Heteroptera. Vol. I.| 1108 581 105 39 903 xx & 462 1880-1893
47. ” ” Vol. IL 607 289 30 22 013. xvi & 416 | 1897-1901
48. Rhynchota Homoptera. Vol. I...) 3238 208 25 13 267 x & 147 1881-1905
- ” Vol. II.
49 f part 1 659 385 42 21 494 xii & 339 1894-1909
yo, Vole HI. ;
L part 2} 161 11 2 Le ae ii & 37 ad
Neuroptera: Ephemeride 32 8 . 1 9) > -
50. P : Ollonata eee eeeees 276 7 1 10 169 | | *** & 420 4) 1901-1908
51. Orthoptera. Vol. I. ... 633 224. 14 22 310*| x & 458 1893-1899 T
02. ” Vol. Il. .... 0. eee 663 138 26 8 120* | viii & 412 | 1900-1909
Total 38637 | 19067 | 1373 1173 18051 | .......- 1879-1915 ©
Botany.
538. Vol, Te cece eee eee ee eee } ( 22 /) Ixviii & 576+) 1879-1888
54. 0g, Le cece cceeceeeceeees L696. | 185 || iv & 576% | 1881-1888
BD. UWL. cesses ee eeseeeuees 7| 11,6265 | “34 |? iv & 711 | 1882-1888
56 gy LV cee ee ) N 5 |) iv & 498 1886-1888
5 a . . 111 144 Vili 1879-1888
ARCHEOLOGY.
Vol. I. (text) ........ cece eee ( viii & 69 | 1888-1902
58 »» IL. (text)... 6... cee eee eee | iv & 47 1899-1902
, » LIT. (text). ...... ce ee eee. iv & 50 1895-1902
PTV. (text) ...scs cece esse. | iv &38 | 1896-1902
59. Vol. I. (plates)............060. 119 iv 1889-1902
60. » Il. (plates).......... 0.06. 98 lv 1899-1902
61. ,, ILL. (plates)............ 000. 82 iv 1895-1902
62. » LV. (plates)... 0.0.6.2 eee 93 iV 1896-1899
63. Appendix (text only).........66. xii & 264 § 1897

* Including allied South- or North-American or Antillean forms figured for comparison.
+ The Introduction, including the systematic list of species figured, was not issued till April 1900, when

Vol. II. was commenced.

~ The Index (pp. 577-619 of Vol. I. and pp. 577-621 of Vol. II.) was replaced, on the completion of
Vol. IV., by a fresh one for all four volumes, which is placed at the end of IV.
§ Calendars, pp. 151-264, not paged.

Srx maps, in addition to the eight included in the Introductory Volume, have been issued in various volumes
of the present work : two, showing the “ Distribution of Freshwater Fishes in Mexico and Central America ”
(Pisces, Introduction, 1908) ; one, illustrating the Distribution of pines, coco-palm, etc. (Botany, Vol. V.,
plate 110) ; one, giving the “ Actual Distribution of Mean Annual Temperatures,” illustrating the distribution
of the Odonata (Neuroptera, Introduction, 1908); one of Guatemala and the adjoining countries, showing
the position of the ruins in Mexico, Guatemala, and Honduras, described by Mr. Maudslay (Archeology,
Vol. II., plate 1); and one of Yucatan and the country to the south of it, showing the position of other
places described by the same author (Archeology, Vol. III. plate 1).

re"
On
-~T
—

ORIGIN, ETC., OF THE FAUNA AND FLORA OF
CENTRAL AMERICA.

{Tuer following summary and conclusions on the origin and geographical distribution
of the Mammalia, Reptilia, Batrachia, Pisces, Arachnida, Chilopoda, Diplopoda, and
Prototracheata of Central America, and the faunistic divisions of the region, have been
kindly supplied by Messrs. R. I. Pocock and C. ‘Tate Regan; and Mr. W. B. Hemsley
has given us a similar account of the Flora. These valuable contributions are based
upon our present knowledge of the subject, bringing the account up to date. As
regards the Insecta, which occupy so large a portion of the work, no satisfactory con-
clusions can be formed till they have been more thoroughly collected and studied in
other tropical regions, and perhaps till we know more of them in a fossil state.-—Eb. |

FAUNA.

MAMMALIA.
By R. I. Pococx, F.R.S.

Since the publication of Mr. E. R. Alston’s volume on the Mammalia of Central
America in 1882, great advances have been made in our knowledge of the subject.
The past history of many of the orders and families has been more or less accurately
ascertained by paleontological research, principally in the United States and in the
Argentine; and the modern methods of collecting and preserving existing material
have led to the discovery of Jarge numbers of genera, species, and subspecies, and
in addition have thrown such light upon the vertical and horizontal distribution of
American Mammals, as a whole, that it has become possible to map their zones and
provinces with much greater precision.

This progress in accuracy of information has been accompanied by a gradual change in
the conception of systematic terms, with the result that what our predecessors ignored
as “local varieties” are now regarded as “species ” and, by a logical sequence, the old-
time species are being given the rank of genera. Quite apart, too, from the names
that have been introduced to designate new forms, considerable changes have taken
place in the nomenclature of long-established species and genera. Opinions may differ
as to the advisability and advantage of the alterations coming under these headings, but
they have to be admitted and dealt with by faunistic and systematic workers.

88 ORIGIN, ETC., OF THE FAUNA.

In view of the additions and modifications thus briefly referred to, it is necessary to
gather together the principal records that have been published during the last quarter
of a century concerning the Mammals of Central America*. ‘The plan adopted will
be to treat the orders separately and to append to each a brief account of its paleon-
tological history and such particulars of its distribution as have a bearing on the
matter in hand.

Order PRIMATES.

The American Monkeys (Cebidee) and Marmosets (Callitrichide) constitute the
Platyrhine tribe of Primates as opposed to the Catarhine tribe embracing the
Monkeys, Apes, and Men of the Old World. ‘The Central American species belong
to genera inhabiting also South America, where they are restricted to the forested
districts east of the Andes, most of them being of tolerably wide range within those
limits. In Central America they are similarly limited to the forest. Of the Cebide,
Squirrel Monkeys (Saimiri=Chrysothrix) extend to Costa Rica, Spider-Monkeys
(Ateles), and Howlers (Alowatta) to Vera Cruz in Mexico, Capuchins (Cebus) to
Nicaragua, and Douroucoulis (Aotus=Nyctipithecus) to Costa Rica and Nicaragua,
but the latter locality needs confirmation. Of the Callitrichide, Geoffroy’s Tamarin
(Edipomidas geoffroy’) ranges from Colombia to Costa Rica.

There are no Monkeys in the Antilles, apart from introduced species, and none in
North America.

There isa good deal of evidence that Monkeys are the descendants of Lemurs, which
date back to the Lower and Middle Eocene of North America and Europe, and survive
at the present time in Africa, Madagascar, and Southern Asia. Oddly enough, this
group of Primates never seems to have entered South or Central America, and no
fossil remains of Monkeys have been found in North America. Nevertheless, Platy-
rhine Monkeys go back to the Upper Miocene in South America if the reference of
Homunculus to that group be, as it appears to be, correct, and no extinct members
of the group (which is, as a whole, more Lemuroid than the Old World Catarhini)
have been found outside that continent. There are one or two facts which suggest
Africa as their original home.

In Madagascar there has been found an extinct Lemur (Archwolemur) which. is
claimed to have Platyrhine characters in its jaws and teeth, and the Oligocene of Egypt
has yielded a Monkey (Parapithecus) structurally bridging the interval between existing
Lemuroids of the Tarsioid (Zarsius) group and the Simiid or monkey-like Primates. If
the interpretations put upon these fossils be correct, it seems that the transitional stages
between the Lemurs and the Monkeys were probably evolved in an Afro-Mascarene

* It is perhaps necessary to explain in this connection that here and elsewhere in this article the term
“ Central America” is used comprehensively, as it is used in the title of the ‘ Biologia,’ to embrace the area lying
between, and including, Mexico and Panama, Lower California being excluded.

MAMMALIA. 89

continent. Coupling this inference with the absence of extinct Monkeys in North
America and of Lemurs in South America, the view that the ancestors of the Platy-
rhini entered South America from Africa is at all events provisionally defensible.

Order INSECTIVORA.

America is not rich in members of this order. Two families, the Talpide (Moles)
and Soricide (Shrews), occur in North America. Although the Talpide range as far
south as the Southern States, they do not appear to enter Mexico or Central America.
The Soricide, on the contrary, are represented in Central America by the three genera
Sorex, Cryptotis (sometimes regarded as a subgenus of the North American Blarina),
and Notiosorer. Of these, Sorex has a wide range over Europe, Asia, and North
America, and passes at high altitudes through Mexico to Guatemala; Notiosorex
inhabits the Southern States of North America (Texas etc.) and Mexico (Sinaloa,
Jalisco). Cryptotis, which replaces in Central America the North American genus
Blarina*, ranges mostly at high altitudes through Central America (Mexico, Yucatan,
Guatemala, Nicaragua, Costa Rica) to Colombia, Venezuela, and Guiana.

Shrews related to Sorex and Blarina have been recorded from the Oligocene of
Europe and North America, and Sores itself goes back to deposits of that age in Europe,
appearing in North America in the Pleistocene, and since no Insectivores of this group
occur in South American deposits f, it seems clear that the Central American species
are southern migrants from North America.

Order CHIROPTERA.

The Central American genera of this order, fifty or more in number, are assigned
to eight families—the Emballonuride, Noctilionide, Phyllostomide, Desmodontide,
Natalide, ‘Thyropteride, Vespertilionide, and Molosside {. Geographically these
families may be referred to two categories: the first containing the Vespertilionide,
Molosside, and Emballonuride, which are represented in the Old World as well as in
America ; and the second containing the remaining five, which are mostiy restricted to
Central and South America, a few genera only occurring in the Southern States of
North America. Of this second category, by far the most important numerically are
the Phyllostomide, the American Leaf-nosed Bats §, the genera of which, occurring in

* A species of Blarina has been recorded from Costa Rica, but there seems to be some doubt about the
correctness of the locality.

+ The only extinct South American Insectivore known is Necrolestes of Upper Miocene age, the nearest
ally of which appears to be the Golden Mole (Chrysochloris) of South Africa,

+ Pending the completion of Dr. Knud Andersen’s ‘ Monograph of the Chiroptera,’ in course of publication
by the Trustees of the British Museum, the classification and nomenclature here followed are those of Miller
(Bull. U.S. Nat. Mus. lvii. 1907). .

§ The Leaf-nosed Bats of the families Rhinolophide, Hipposideride, and Megadermide are restricted to
the Old World. .

BIOL. CENTR.-AMER., Introd. Vol., June 1915. N

90 ORIGIN, ETC., OF THE FAUNA.

Central America, are about equal numerically to the genera of all the other faim ilies
put together.

A few only of the more important genera need be mentioned.

‘The Vespertilionide are cosmopolitan to the limit of tree-growth. Three familiar
Old World forms occurring in our area are: Myotis, extending from North America to
the Argentine; Eptesicus, spreading as far south in America as Guatemala; and L%pi-
strellus, reaching South Mexico. Corynorhinus, the American representative of the
Old World Plecotus, ranges from North America into Mexico. Other American
genera are: Dasypterus, extending from North to South America; Rhogoeéssa, from
Central Mexico to Venezuela; and Beodon, known only from Jalisco, in Mexico.
Finally, Antrozous, formerly, but wrongly, affiliated with Plecotus and ranging from the
Southern States of the Union to Central Mexico, constitutes with the Australasian
genus Nyctophilus a special subfamily of the Vespertilionide.

The Molosside are more southern in distribution than the Vespertilionide. In the
Old World their northern limit is South Europe and Asia. One genus only, Nyctinomus,
found also in Europe, Africa, Asia, and Australia, reaches America, where it spreads
from the Southern United States, though Central America, to Chili. Of exclusively
American genera, Promops and Molossus pass from Mexico into South America, while
Eumops has a northern range into the Southern States.

Of the Emballonuride, mainly a tropical family, not one of the Old World genera
occurs in America, and the American genera (e.g. Saccopteryx, Peropteryx, and
Diclidurus) extend from Central into South America.

Of the families confined to America, the Noctilionide contains two genera—one South
American, the other (Nocti/io) passing northwards to South Mexico; the Natalide and
Thyropteride contain one genus each, namely Natalus and Thyroptera, ranging
respectively from South America to Central Mexico and Honduras; the Phyllostomide,
divisible into several subfamilies, have, broadly speaking, a distribution similar to that
of the other exclusively American families just mentioned, though some of the genera
have a wider, others a narrower, range. For example, of the Chilonycterine, Mormoops
reaches Texas, Chilonycteris and Pteronotus do not go north of Mexico; of the
Phyllostomine Xenoctenus is only known from Costa Rica, Glyphonycteris passes
southwards from that country to Peru, whereas Otopterus extends from the Southern
States of North America to Guatemala; of the Glossophagine, Cheronycteris reaches
from South America to Arizona. The single species of Halonycteris is known only from
Costa Rica, while Lichonycteris, also monotypical, has been recorded from N icaragua
and Guiana, Similarly, in the Stenodermine, the monotypical Centurio and Ectophylla
are Central American, but the other Central American genera range into tropical
South America, Finally, the family Desmodontide, or true Vampyres, differing from
the Phyllostomide by their large shear-like anterior teeth, is represented by two

genera, Desmodus and Diphyllus, which are distributed from Southern Mexico to
Paraguay and Brazil.

MAMMALIA. . 91

The small number of extinct Chiroptera known, 7.e. a few genera belonging to the
Hipposideride and Vespertilionide from Eocene and Oligocene deposits of Europe,
throw no light upon the present distribution of the order.

Order RODENTIA.

In the number of species and genera inhabiting Central America, this order rivals,
if it does not surpass, the Chiroptera, the four usually admitted suborders (Sciuro-
morpha, Myomorpha, Hystricomorpha, and Lagomorpha *) being well represented.

Suborder SCIUROMORPIIA.,

Two families of this suborder have to be considered, namely the Petauristide or
flying Squirrels and the Sciuride or true Squirrels, Ground-Squirrels, and Marmots.
The one American genus of Petauristide, Sciwrepterus, ranges from Europe and
Asia into North America, where it extends from Alaska to Guatemala. Of
the Sciuride, the Chipping Squirrel (Ewtamias) and the Souslik (Citel/us) have a
distribution as extensive as that of Sciwropterus, whereas an allied form, Callospermo-
philus, and the Prairie Marmot ( Cynonys) are restricted to the South and Western
States of North America and North Mexico. ‘True arboreal Squirrels are represented
by vast numbers of species and subspecies in North, Central, and South America, as
well as in Europe, Asia, and Africa; but the genera and subgenera to which they have
been referred require revision and re-definition before their mutual affinities can be
accurately ascertained. Of the Central American genera, Sciurus (s. s.) appears to
ve the only one that occurs in the Old World, where it ranges from Japan to Ireland.
In America it is generally distributed throughout the States, occurs everywhere in
Central America at high and low levels, and enters the northern countries of South
America (Colombia, Ecuador). An allied form with many species, Guerlinguetus
(Parasciurus), is credited with a range from the United States to Peru. Others that
may be mentioned are Baiosciurus (extending from Mexico to Nicaragua), Syntheo-
sciurus (known only from Chiriqui in Panama), and Mrcrosciurus (alleged to spread
from Costa Rica to Peru f).

Genera regarded as primitive Sciuromorphs were abundant in North America
(Ischyromyide) and Europe (Pseudosciuride) during early Tertiary times (Hocene and
Oligocene), extinct Marmots (Palearctomys) and the still existing Cynomys have been
traced back to the Upper Miocene in North America, while Sciurus itself has survived
in Europe and North America since the Upper Oligocene. Since no extinct forms are
: * Apart from the Lagomorpha, these suborders are not perhaps susceptible of precise definition, They

are here maintained for the sake of convenience.
+ Beavers (Castoride) should perhaps be added to this section, since they range in North America at least
as far south as Sonora. In the Old World, Beavers occur in Northern Asia and North and Central Europe.
NZ

92 ORIGIN, ETC., OF THE FAUNA.

known in South America, unless of quite recent date, it seems clear that Central
America received this prominent element of its fauna from North America. :

Beavers also seem to have originated in North America, where they date back to
the Middle Oligocene, but Castor itself appeared in Europe in the Pliocene and in
North America in the Pleistocene.

Suborder MYOMORPHA.

- The Central American Myomorphs belong to the three families Muride, Hetero-
myide, and Geomyide, the two last-named being restricted to America.

The genera of Muride belong to the three subfamilies Microtine, Neotomine, and
Cricetine. Ofthe Microtine, Microtus (s. s.) extends over the greater part of Europe,
Asia, and North America, and enters Mexico (Puebla, Jalisco, Oaxaca). Two sub-
genera are known only from Central America, namely, Orthriomys from Oaxaca and
Herpetomys from Guatemala, both at high altitudes. The other genus Pitymys has a
singular distribution, ranging in the Old World from South and Central Europe to
Asia Minor (the Mediterranean Area), and through the Eastern States of North
America to Mexico (Vera Cruz).

Of the Neotomine, Xenomys, Hodomys, Nelsonia, and Teanopus are confined to
Mexico, while Neotoma, represented by a host of species, overlaps them to the north
and south with a range from the Southern States of North America to Guatemala.

The Cricetinz have an extraordinarily wide distribution, namely Africa, including
Madagascar, Europe, Asia, and practically the whole of America. The Central
American genera are too numerous to mention in detail. Some occur in North and
Central America, others in Central America alone, others are common to Central and
South America, while others extend over the three areas mentioned. To the last
category belong Reithrodontomys, Oryzomys, and Sigmodon, ranging from the Southern
States of the Union to Ecuador, Chili, and Peru, respectively. As examples of the
first group may be cited Peromyscus (Hesperomys) distributed from Labrador to
Panama, and Baiomys and Onychomys from the Southern States to Guatemala and
Panama. Peculiar to Central America are Nyctomys and Ototylomys, ranging from
Mexico to Panama and Guatemala, and Sigmodontomys known only from Costa Rica,
while Akodon, extending all over South America from Costa Rica, and Nectomys,
occurring in Nicaragua, Ecuador, and Colombia, are representatives of genera common
to Central and South America.

The genera of Heteromyide belong to two subfamilies, the Heteromyine and
Dipodomyine. Of the latter, Perodipus and Dipodomys (Jerboa-Mice) are restricted
to the South and South-western States of North America and to Mexico (Zacatecas,
Vera Cruz, etc.). Of the former, Heteromys, Perognathus, Liomys, and Chetodipus
occur in the Southern United States and Mexico, the first passing southwards into

MAMMALIA. 93

Colombia and Venezuela. The Geomyide, or Pocket-Gophers, are mainly Central
American, Thomomys, Cratogeomys, and Geomys ranging into the Southern States of
the Union, Platygeomys, Zygogeomys, etc., being Mexican, Orthogeomys extending
from Mexico to Guatemala, and Macrogeomys occurring from Nicaragua to Panama.

The paleontological history of the Voles (Microtine) before the Pleistocene is some-
what meagre, but Microtus has been traced back to the Pliocene in Europe and to the
Pleistocene in North America. Records of the Cricetine are more complete, and
extend back to the Lower, Middle, and Upper Oligocene in Europe and North America,
the existing Peromyscus dating in North America to the Miocene. In South America
Cricetines appear to be of recent (Pleistocene) date. The Heteromyide and Geo-
myide, closely related families, seem to have originated in North America, where
they have been traced respectively to the Lower and Upper Oligocene.

The palzontological evidence, therefore, suggests that the ancestors of the Central
and South American Myomorpha came down from the north, This is no doubt true
of the Geomyide and Heteromyide, but it is not so certain of all the Cricetines. The
failure to discover fossil remains of small rodents such as these in mid-Tertiary
(Upper Miocene) deposits in South America cannot be regarded as conclusive evidence
of their absence there at the time the deposits were laid down; moreover, the
occurrence of genera of this subfamily in Madagascar strongly suggests their existence
in Africa at a sufficiently early date to permit their migration thence into South
America by the land-connection which there are other grounds for thinking may have
joined these two continents together.

Suborder HYSTRICOMORPHA.

Three families—the Octodontide, Dasyproctide, and Hystricidee—occur in Central
America, but neither is peculiar to the country, though the Dasyproctide are found
elsewhere only in South America and the West Indies. None of the Central
American genera of these families enters the United States.

Of the Dasyproctide, Dasyprocta (Agouti) ranges from Brazil through Panama to
Mexico, probably Vera Cruz; <Agouti or Calogenys (Sooty Paca) also passes from
Panama to Vera Cruz. ‘The Hystricide are represented by Coendou (= Synetheres),
the common South American Tree-Porcupine, which extends from Bolivia and Brazil
to the mountains of Mexico. In North America its place is taken by Lrethizon,
distributed from Alaska and Canada to Arizona. One other genus of this family,
Chetomys, occurs in South America only. ‘These arboreal Porcupines constitute the
subfamily Erethizontine as opposed to the Hystricine or Ground-Porcupines of the
Old World. ‘The Octodontide, generally distributed throughout South America,
comprises a large number of genera, two of which, Loncheres and Proechimys, enter.
Central America, the latter as far as Nicaragua, the former only to Panama. A third,
Hoplomys, occurs in both these countries.

94. ORIGIN, ETC., OF THE FAUNA.

Although South America must be regarded as the present headquarters of the
Hystricomorphs, on account of the number and diversity of the genera that occur there,
the group is not restricted to America. There are several African genera belonging
to or closely allied to the Octodontide ; and Porcupines (Hystria, Atherura) are found
in Tropical Asia as well as Africa.

The paleontological history of the Hystricomorphs is very different from that of
the other suborders of Rodentia. Genera regarded as ancestral members of the group,
and constituting the family Theridomyide, were living in Europe from the Middle
Eocene to the Upper Oligocene, and true Porcupines, related to the existing species of
Hystrix, were in Europe in the Middle Miocene; but none of these extended into
North America. On the other hand, genera assignable to the Octodontide and
Erethizontine Porcupines suddenly appear in South America in Upper Miocene
deposits—that is to say, considerably later than the date of the first known appearance
of the group in Europe. The available evidence therefore points to the conclusion
that this suborder made its way into South America from Africa, and subsequently
spread northward into Central America and into North America, Erethizon dating
from the Pleistocene in the latter country.

Suborder LAGOMORPHA.

The Ochotonide (Picas)—which occur in Central Asia and spread through Western
North. America from Alaska to California, Utah, and Montana—do not enter Mexico.
‘The Leporide have a much wider range. Three genera have been recorded from
Central America, namely Lepus, Sylvilagus, and Romerolagus. ‘The last is mono-
typical, and has been discovered at Puebla, Popocatepetl, etc., in Mexico at a height
of 10,000 to 12,000 feet. Sylvilagus runs right through Central America from the
Southern United States into South America. Jepus, in the modern sense, on the other
hand, does not go south of Mexico (Durango, Tamaulipas, Potosi, etc.), whence it
spreads northwards to Greenland and is also found in Asia, Europe, and Africa,

The earliest paleontological records of Hares (Leporide) are from the Lower
Oligocene of North America (Palwolagus). Lepus itself was in existence in the Upper
Oligocene of that continent, and appeared in Europe in the Lower Pliocene. It seems
therefore that this suborder of Rodents was evolved in North America, and thence
made its way into the Old World and into Central and South America.

Order CARNIVORA.

_ The terrestrial or fissiped Carnivores of Central America belong to the five families
Canide, Procyonide, Urside, Mustelide, and Felide.

Canide:—The genus Canis is represented in North America by many species or
local races belonging to groups typified by C. occidentalis and C. latrans, corresponding

MAMMALIA, 95

respectively to the Wolves and Jackals of the Old World. Both groups occur on
the Mexican plateau—distinct species of the latrans or Prairie Wolf type having
been recorded from Tamaulipas, Durango, Puebla, etc., as far as Guatemala, and an
unknown form is said to occur in Costa Rica. A second genus, Urocyon, commonly
regarded as a Fox, extends from the Eastern States of North America, through
Central America (Mexico, Yucatan, Guatemala), to Colombia in South America.

There are reasons for thinking that Urocyon is more nearly allied to the South
American group of Canide typitied by Cerdocyon thous than to Vulpes. If this be so,
the genus constitutes an interesting geographical link between the Canide of North
America and those of South America. In the iatter continent there is a very large
number of species ranging from the extreme north to Cape Hern and the Falkland
Islands. These are referable to several groups possibly of generic status; but none
of these can be definitely affiliated with Canis or Vulpes or other genera inhabiting
North America and the Old World. The apparent absence from the tropical countries
and highlands of Central America of Canide related to the forest and Andean species
of South America is a singular fact.

Mustelide.—The dominant Central American Mustelids appear to be Skunks, which
are represented by Mephitis ranging from Canada to Guatemala, Spilogale from the
Southern United States through Mexico to Costa Rica, and Conepatus with much the
same northern limits as Spiloga/e, but extending throughout South America to ‘Tierra
del Fuego, mostly at high levels. Weasels (Mustela) range all over Europe, Central
and Northern Asia, North America, and thence southwards through Central America
into the northern countries of South America. The remaining terrestrial forms, the
Tayra (Zayra = Galera) and the Grison (Grison = Galictis), are more southern in
distribution. The former is found in Vera Cruz, Nicaragua, Panama, Guiana, Peru,
Brazil, etc., and the latter in Honduras, Yucatan, Costa Rica, and thence southwards to
Patagonia. Finally, Otters (Lutra) are represented in Cental America by two species,
one described from Jalisco, the other from Nicaragua. This genus is practically
cosmopolitan in the Old World, apart from Madagascar and the Australian Region,
its exceptionally wide range being possibly connected with independence of
terrestrial barriers resulting from its power of making its way by swimming along
coast-lines.

Felide.—Cats are abundant everywhere, and belong to many distinct groups, probably
of generic value. ‘The short-tailed or Lynx-group, which is widely distributed over
Northern Europe, Asia, and North America, enters Mexico (Tamaulipas, Sinaloa). The.
Puma-group, which has no close kinship with Lions and Leopards of the Old World,
ranges from the confines of Canada to Patagonia and occurs in Central America at both
high and low levels from Mexico to Panama. The other species of this family—
belonging to a variety of groups akin to the Jaguar (F. onca), Ocelot (#. pardalis),
Eyra (Ff. eyra) groups, etc.—are related to the South American species of those

96 ORIGIN, ETC., OF THE FAUNA.

names. Both Ocelots and Jaguars frequent the forested districts of South America,
and pass through Central America into the United States north of Mexico. There is
no doubt that the Jaguar is closely allied to the Leopards, and is the only American
member of the genus which comprises also the Lion, Tiger, Leopard, and Ounce
of the Old World. Ocelots, too, are not apparently generically separable from
several of the Eastern Asiatic species, and there appears to be a form closely akin to
them in tropical West Africa.

Procyonide.—The five admitted American genera of this family occur in Central
America, which in this respect is richer than either the northern or southern moieties
of the New World. Procyon (Raccoon) has a range almost equal to that of the Puma
in its extension from near the borders of Canada to Paraguay. Subspecies akin to
the typical North American form (dotor) reach Southern Mexico; two species appear
to be restricted respectively to Yucatan and Panama, while the Scuth American
P. cancrivorus enters the latter State. A peculiar species of MNaswa (Coaiti Mondi)
occurs on Cozumel Island, Yucatan, the rest of the Central American forms recorded
from Mexico (Colima, Chihuahua), Yucatan, Guatemala, and Panama being regarded as
subspecies of the South American WV. narica; Bassariscus (Cacomistl) ranges from
Texas and Oregon through Mexico to Guatemala and Panama; Bassaricyon is
represented by species from Nicaragua, Costa Rica, and Panama, and occurs as far
south as Ecuador; while Potos, better known as Cercoleptes (Kinkajou), ranges
from Mexico (Vera Cruz) Costa Rica, and Guatemala into Colombia, Ecuador,
Venezuela, etc.

In the Old World the family is represented by lurus (Panda) of the Eastern
Himalayas and probably by luropus of Moupin in Eastern Tibet. The latter,
however, is claimed by some authorities to be Ursine in its affinities. It shares the
characters of the Urside and Procyonide, and may perhaps be regarded as the living
link between the two families.

Urside.—Bears akin to the North American Black Bear (Ursus (Euarctos) ameri-
canus) extend into Mexico as far as Coahuila and Chihuahua. South of Mexico the
family is unknown in Central America; but it reappears in the northern Andean
district of South America, where a genus (Zremarctos), distinct from Ursus, is now
found—and this genus, oddly enough, also appears to contain the Central Asiatic
Black or Tibetan Bear, which ranges from Baluchistan through the Himalayas to
Japan.

The paleontological history of the families of Central American Carnivora need not
be discussed in detail. The evidence, incomplete in details though it be, shows that
they are without exception descended from northern forms that inhabited North America
or Europe or both continents during Middle or early Tertiary times, the Canide dating
back to the Upper Eocene, the Procyonide to the Lower Miocene in North America,
the Urside to the European Upper Miocene where they apparently blend with the

MAMMALIA. 97

Canide, the Mustelide to the Lower Oligocene, and the Felide *, even Felis itself, to
the Middle Miocene of Europe and North America. No extinct forms so early in
time have yet been discovered in South America. Hence it is needless to look
beyond North America for the immediate origin of the Central American and South
American forms. But no data, paleontological or otherwise, seem to have been
discovered as yet to explain the almost complete absence of Canide from the southern
countries of Central America and the discontinuity in the distribution of the Bears
above alluded to. Nor in the case of such genera as Galera, Grison, Nasua, and others
is it clear whether their presence in Central America is due to northward migration
from South America, or whether it is attributable to their settlement in the former
area on their way to the southern continent.

Order ARTIODACTYLA.

The Bunodont or non-ruminant Artiodactyls are represented in America by the
family Tayassuide (formerly Dicotylide), commonly known as Peccaries. ‘The
described species may be referred to two categories, regarded sometimes as genera,
exemplified by the Collared Peccary (Tayassu tajacu = torquatus) and the White-
lipped Peccary (Z. pecari = labiatus). Species of the former category range from
Texas (Guadalupe) to the Argentine, and have been recorded from Mexico (Sonora,
Colima, Puebla), Guatemala, and Panama ; those of the second category spread from
Campeche to Paraguay.

Three families of ruminant Artiodactyls inhabit Central America, namely the
Bovide, Antilocapride, and Cervidee. The first two, represented by the Bison (Bison),
the Rocky Mountain Sheep (Ovis), and the Prongbuck (Antilocapra), are essentially
North American. ‘The Bison inhabited Mexico in historic times. The Sheep and the
Prongbuck penetrate Mexico as far as Chihuahua. The Cervide or Deer belong to
two genera—Odocoileus, which extends from the United States, through Mexico to
Panama, and thence into South America, and Mazama, ranging from Mexico (Vera
Cruz) and Yucatan to Brazil and the Northern Argentine.

The paleontological history of the Peccaries is fairly well known, many genera of
the family having been discovered in the North American Tertiaries, dating from the
Lower Oligocene. But our knowledge of the descent of the ruminant Artiodactyla is
singularly meagre in the matter of details. ‘The Central American Cervide (Odocotleus,
Mazama) belong to a group of the family (Telemetacarpalia) which, with a few
exceptions, is confined to America. Possibly they are descended from a form
(Blastomeryx) which existed in North America in the Lower Miocene. Odocorleus
itself goes back to the North American Pliocene, but it seems difficult to avoid the

* Excluding the extinct Oligocene Macherodonts, which have been recently claimed as theirancestors. .

BIOL. CENTR.-AMER., Introd. Vol., June 1915. ©)

98 ORIGIN, ETC., OF THE FAUNA.

conclusion that Mazama, typically a South American genus, is a more primitive type,
unless it be degenerate, in its simple one-spiked antlers. Nevertheless, the absence of
extinct Deer in South America, except in late Tertiary (Pleistocene) deposits, precludes
at present the view that the family was evolved in that country and subsequently made
its way northwards.

The Antilocapride are not certainly known from deposits in North America older
than the Pleistocene, and none have been discovered elsewhere. Possibly the North
American mid-Miocene genus Dromomeryx was the ancestral form.

The oldest-known forms of Caprine (Ovis) and Bovine (Bison) are European.
The Bovinze do not appear to be definably distinct from the so-called Tragelaphine
Antelopes, which in Europe go back to the Upper Miocene. A Caprine (Cridotherium)
is of that date also. Hence it appears that the Artiodactyle Ungulates of Central
America must be regarded as of northern descent.

Order PERISSODACTYLA.

Of the existing families of this order only the Tapiride occur in Central America,
where they are represented by two species of the genus Tapirella (Elasmognathus),
namely bairdi, ranging from South Mexico to Panama, and dow?, from Guatemala to
Nicaragua and Qosta Rica. The Tapirs of South America, where they extend from
Venezuela to the Northern Argentine, are referred to a distinct genus, Zapirus.
Outside Tropical America the family contains but one representative, namely, Rhzno-
chorus indicus, from the Malay Peninsula and Sumatra.

Tapirs referred to a variety of genera inhabited North America from the Lower
Eocene and Europe from the Lower Oligocene. ‘There are gaps in the genealogy of
the recent forms, but species akin to them have been discovered in Pliocene deposits in
Europe and Asia and possibly in contemporaneous bedsin North America. From this
it seems tolerably clear that the existing genera are the survivors in the East Indies
and in Tropical America of a family formerly widely distributed in the Northern
Hemisphere.

Order SIRENIA.

One genus of this order, the Manatee (Zrichechus = Manatus), occurs in the rivers
of eastern Central America and South America. It also inhabits some of the rivers of
Western Africa, but is not found elsewhere.

‘There is evidence that the Sirenia were evolved in North Africa from the stock that
also gave rise to the Proboscidea. Genera of Eocene age have been discovered in
Egypt and also in the West Indies. Although Zrichecus itself is only known as an
existing genus, its presence in rivers and estuaries on the eastern and western shores
of the Atlantic and its avoidance of the open sea have been cited as evidence of a
continuous coast-line between Africa and Tropical America.

MAMMALIA. 99

Order EDENTATA.

Eliminating the Old World genera Orycteropus and Manis, this order is confined to
South and Central America. Cholepus and Bradypus, the two genera of Sloths or
Bradypodide, pass from South into Central America, the former as far as Costa Rica and
the latter to Nicaragua. Myrmecophaga and Cyclopes, two of the Anteaters (Myrme-
cophagide), also go as far as Guatemala; the third genus, Tamanduas, extends
northward to Tabasco and Vera Cruz in Mexico. Of the large number of South
American genera of Armadillos (Dasypodide) two only enter Central America, namely
Cabassous (Lysiurus), which ranges from Brazil to Honduras, and Dasypus (= Tatu or
Tatusia), which overlaps it both to the north and south, with au extension from the
Argentine to Texas, just touching the latter state at Brownsville.

The origin and centre of evolution of the Edentata is at present unknown. The
only extinct forms that have been claimed to be their ancestors are the Txniodonts or
Ganodonts of the Eocene of North America. Since this view has been authoritatively
disputed, it needs no further consideration here. Of more importance, from our
present point of view, is the mid-Eocene North American genus Metachetromys, believed
by some to be a primitive Armadillo. It is true that the Armadillos (Dasypodide) are
the least specialised of all undoubted Edentates, living or extinct, and no reason can
be alleged against the likelihood of the discovery of an early type in the Eocene of
North America, but since Metacheiromys unfortunately has few, if any, of the
characters distinctive of Edentates as we know them, it cannot be maintained as
established that the order is of North American origin. It may be suggested that
their ancestors entered South America from Africa. However that may be, the
~ evidence of fossil forms shows that South America has been the centre of evolution
of the large number of existing and extinct genera that have been discovered. In
South America, Armadillos (Dasypodide) date back to the Upper Miocene, the Dasy-
podine (olim Tatusiine) themselves, which are now found in Central and North
America, ranging from that date to the Pleistocene. Hence the presence of Dasypus
in Central America and Texas must be assigned to northward migration. And the
same must be claimed of the Sloths (Bradypodide) and Anteaters (Myrmecophagide).
No extinct forms of these families have yet been discovered, but abundant remains of a
wholly extinct group, intermediate in many respects between the two, but less specialised
than either, have been discovered in South American Miocene and later deposits.
These are the Ground-Sloths (Mylodontide and Megatheriide), which at a later date
(Pliocene and Pleistocene) lived also in North America.

Order MARSUPIALIA.

The American Marsupials belong to two families, the Didelphyide (or Opossnms) and

the Cenolestide. ‘The latter, represented by the two existing species of Canolestes
0 2

100 ORIGIN, ETC., OF THE FAUNA.

from Ecuador and Colombia, does not appear to reach Central America; but no fewer
than five genera of the former occur in our area, namely Didelphys, Metachirus,
Caluromys (=Philander), Marmosa, and Chironectes. Of these Didelphys is the only
one that is found in North America, where it ranges from about the 40th parallel
of N. lat. to Texas, thence passing southwards through Mexico, Nicaragua, Yucatan,
Guatemala, and Panama as far as the Argentine and Chili. The others are Central
and South American, Marmosa ranging from Mexico (Oaxaca), Costa Rica, and Panama
to Chili, Wfetachirus trom Mexico (Vera Cruz), Guatemala, and Costa Rica to the
Argentine, Caluromys from Mexico (Tabasco) to Paraguay, and Chironectes, the
aberrant water-opossum, from Guatemala to Brazil.

Extinct Marsupials belonging to the Didelphyide, and closely related to the existing
genera of that family, have been discovered in Lower Eocene deposits in North
America and as late as the Lower Miocene in Europe, and in what are believed to be
Upper Cretaceous beds in South America, Formerly, therefore, the family was very
widely distributed. Nevertheless, Opossums appear to have survived uninterruptedly in
South America, at all events since the earliest Tertiary times, and that country is now
their headquarters. These facts, coupled with the absence of paleontological evidence
that the group survived in North America after the Oligoceue and in Europe after the
Miocene, justify the supposition that the existing genera are recent immigrants from
South into Central and North America.

SUMMARY AND CONCLUSION.
ORIGINS OF THE CENTRAL AMERICAN MAMMALIAN FatNa.

The foregoing account shows that the Mammalian fauna of Central America is a
mixture of two elements, namely, of forms which are dominant in North and South
America respectively ; and the paleontological history, during Tertiary times, of the
orders concerned, supplies—in part, at all events—the explanation of the intermixture. |
So far as this history has been read, it furnishes strong evidence that the Mammals
may be referred to two categories. To the first belong the Insectivora, Carnivora,
Artiodactyla, Perissodactyla, and the Rodentia (with the exception of the Hystrico-
morphs and possibly some of the Cricetine Myomorphs), which were evolved
through long ages in the Northern Hemisphere and inferentially passed from North
into South America by way of Mexico and Panama. ‘The second comprises the
Edentates, Primates, Marsupials, and the Hystricomorphous Rodents which, from
whatever country they may originally have come, have undergone a prolonged course
of evolution in South America and migrated thence into Central or even North
America. ‘That is to say, whereas North and South America have been independent
geographical centres for the evolution of Mammals in the Western Hemisphere, the
part played in the main by Central America has been that of a bridge joining these
centres together and permitting the intermingling of the independently developed

MAMMALIA. 101

faunas. A few genera of Chiroptera and Rodentia restricted, so far as is known, to
Central America, may have been evolved within the area; but these do not alter the
general character of the country as a faunistically transitional tract between the
northern and southern portions of the Western Hemisphere. |

That the intermingling and cross-migrations, which give the stamp to the existing
Central American Mammalian fauna, began in the Miocene and have continued
uninterruptedly since that date may be inferred from paleontological and geological
data, which support the conclusion that the Northern and Southern Americas, separated
by sea over what is now the isthmus of Panama from the earliest Eocene (Paleocene)
to the end of the Oligocene, were finally joined by the elevation of that isthmus during
the Miocene Period.

Possibly, as held by some authorities, Central America to the north of Costa Rica
and Panama was temporarily connected with South America by way of the Greater
and Lesser Antilles in the Early Oligocene; and possibly there was a stiil earlier union
during the Cretaceous of the western portions of North, Central, and South America.
However that may be, it seems tolerably certain that the main streams of migration
passed by way of Costa Rica and Panama in comparatively recent Tertiary times.

Reference has more than once been made to the possibility of South America
having received the ancestors of some of its characteristic forms of Mammals (Platy-
rhine Monkeys, Hystricomorph Rodents, Sirenians) from Africa by a direct trans-
Atlantic bridge between the two continents. Other groups of animals supply evidence
for the existence of this union. But if the mammals made use of it, it must have
endured into early Tertiary times. This, however, has been disputed.

Another point, also under discussion, may here be referred to, although it has no
direct bearing, so far as is known, upon the fauna of Central America. This is the
possibility of a direct connection between South America and Australia. Perhaps the
most cogent evidence for this is supplied by certain genera of invertebrates, probably
in great part ancient types. So far as Mammals are concerned, the evidence rests
upon the claimed relationship between the so-called Tasmanian or Marsupial Wolf
(Lhylacinus) and some extinct (Upper Miocene) Sparassodont Mammals of South
America, coupled with the later date of fossil Marsupials in Australia, the absence of
their remains in Tertiary deposits in China, and the absence of living forms in South-
east Asia to the west of “ Wallace’s Line.” Also there is the undeniable kinship
between the Australian Dasyurids and the American Didelphyide, of which Marmosa
is alleged to be the most primitive type. It would be out of place further to discuss
these questions here; but the facts, as they stand, are suggestive of the origin of the
Australian Marsupials from a South American stock. Moreover, if Thylacinus be of
the same family as the Sparassodont Prothylacinus, it may be held as perhaps
probable that the connecting land-mass permitting the migration persisted into early

‘Tertiary times.

102 ORIGIN, ETC., OF THE FAUNA.

FAvUNISTIC DIVISIONS OF CENTRAL AMERICA.

Central America has been described above as transitional with respect to its
Mammalian fauna between North and South America. ‘This statement, however, is
perhaps too general and apt to give a mistaken impression of the actual facts; for the
transition cannot be described as complete. Its incompleteness, however, is not due
to the existence of any physical barriers to migration, unless temperature and moisture
can be described as such. But, as will appear in the sequel, the faunistic do not
coincide with the political divisions of this land-area.

The central portion of Mexico forms an extensive plateau rising some 9000 feet
above the sea, with mountains nearly twice that altitude. This tableland to the north
is continuous with, and closely resembles in its characters, a great tract of comparatively
dry territory stretching into North America and embracing the southern parts of
California, Lower California, Nevada, Arizona, and nearly the whole of Texas.
Southwards the plateau is continued by the mountain ranges which stretch throughout
the southern States of Central America In Mexico it rises somewhat abruptly
from the lower-lying country towards the coast on each side of it; and this country,
covered for the most part with tropical forest, extends to east and west of the plateau
up to or almost up to the confines of the United States. ‘Tropical forest-conditions
also prevail over the greater part of Central America to the south of the plateau,
although, as has been stated, the country is almost everywhere broken up by
mountains rising several thousand feet above sea-level.

It is needless to recapitulate here the facts set forth above regarding the distribution
of the families and genera of Mammals inhabiting Central America. It is clear, how-
ever, that those believed to have undergone a long course of evolution in South
America (Primates, Edentates, Hystricomorphous Rodents, and Marsupials) are wholly,
or in the main, restricted to the forested tracts above described, although not by any
means occurring throughout such districts. Most of the genera and families do not
pass north of Vera Cruz, in Mexico, and Guatemala. Notable exceptions are the
Armadillo (Dasypus) which reaches Texas, some of the Opossums which occur on the
Mexican plateau—one, indeed, penetrating far to the north in the United States,—and
the Porcupine (Erethizon), belonging to the same family as Coendow and ranging
from Arizona to Alaska.

Similarly, with the groups which, although known to have been evolved in North
America or Eurasia, are now dominant in the southern tropical districts of America,
such as the Tapirs, Peccaries, Brockets (Mazama), most of the Felide, some Procyo-
nid (Nasua, Potos), and Mustelide (Zayra, Grison). These genera, usually regarded
as intruders from the south, though the evidence on that head seems inconclusive, are
mostly restricted to forest-covered tracts of Central America. And the same thing
applies to the Bats of the families Phyllostomide, Desmodontide, ete.

MAMMALIA. 103

On the other hand, there are a great many genera like the Prongbuck (Antzlocapra),
Bison (formerly), Rocky Mountain Sheep, Beaver, Prairie-Marmot (Cynomys), Chipmunk
(Lutamias), Black Bear, Wolf, Lynx, and other typically North American forms which
are restricted in Central America to the Mexican plateau or to the highlands just to
the south of it.

These differences in distribution exist quite apart from the occurrence throughout
Central America of many families and genera like the Hares (Leporide), the typical
Squirrels (Sczwrus etc.), the Shrews (Soricide), Pocket Gophers (Geomyide), Raccoon
(Procyon), Deer (Odocoileus), and others.

Zoogeographers have given practical expression to the facts above set forth by
referring the plateau of Mexico to the same zoological region as the southern States of
North America. Out of a number of titles that have been proposed for this region,
Sonoran is the one preferred. The rest of Central America, on the other hand, is
regarded as a subregion of the Neotropical Region—sometimes called the Neogzic
Realm,—which embraces, in addition, the Antilles and the whole of South America.
On the western side of the plateau this region stretches to about lat. 25° N., and on
the eastern side a little farther, stopping short near the Nueces River in Texas.

The subregional distinctions between the tropical portions of Central America and the
adjacent portions of South America are not sharply defined, and rest upon the not very
satisfactorily established statistics of the comparatively small number of typically
South American genera which occur in those areas of Central America, and the
presence in the latter of certain Sonoran forms (Soricide, Geomyide) whose southward
range practically stops short at Panama.

Similarly, the Antillean subregion is characterised mainly by the poverty of its fauna
in types occurring in Central and South America, It is noticeable that the orders of
known northern origin are practically absent. Some of the Chiroptera (Eptesicus,
Myotis) are, perhaps not surprisingly, exceptional. Most of the genera of this order
belong to the Tropical American family Phyllostomide. For the rest, the Hystrico-
morph Redents are represented by Capromys, Plagiodontia, and Loncheres, all Octodonts,
the first two being peculiar to the subregion, by Dasyprocta, and possibly by a species
of Coendow. Very interesting is the occurrence of a Cricetine Myomorph, Megalomys,
peculiar to the Lesser Antilles. Monkeys, however, are absent, and the Edentates are
represented only by one species of Armadillo (Dasypus) from Grenada. Finally, the
Insectivora are exemplified by the Cuban and Haitian Solenodon, whose nearest living
allies are the Centetide of Madagascar (Centetes etc.) and West Africa (Potamogale).
The family Solenodontide, however, dates back to the Lower Oligocene in North
America, where one genus, MZicropternodus, has been discovered.

Until the Mammalian Paleontology of the West Indies has been worked out, the
geological history of these islands cannot be accurately read. Nevertheless, temporary
union between the Lesser Antilles and South America is indicated by the occurrence

104 ORIGIN, ETC., OF THE FAUNA.

of such genera as Loncheres, Dasyprocta, and Dasypus. Furthermore, the restriction of
Capromys to Cuba, Jamaica, and some of the islands towards Central America and
the kinship between this genus and living and fossil South American genera suggest
a connection between Central America and those islands of the Greater Antilles.
But this conclusion cannot at present be reconciled with the absence of other Central
American forms, both of northern and southern origin, from the Greater Antilles.
If, on the other hand, the Hystricomorphs passed into South America from Africa or
South Europe, and if the West Indies formed part of the transatlantic land, the
faunistic resemblance between Haiti, Cuba, Jamaica, and South America supplied by
Capromys and Plagiodontia may be due to the derivation of the fauna from a common
African source. ‘The relationship between Solenodon of Cuba and Haiti and the
Afro-Mascarene Centetide has an interesting bearing on this question, although, if
Micropternodus be, as alleged, a Selenodont Insectivore, Cuba and Haiti must
presumably have been connected with some part of North America.

PRINCIPAL WORKS CONSULTED.

Auston, E. R.—Biologia Centrali-Americana. Mammalia, 1879-1882.
Gavow, H.—The Wanderings of Animals. Cambridge Manuals of Science and Literature, 1913.
Lypexker, R.—A Geographical History of Mammals. Cambridge Geographical Series, 1896.

Merriam, C. Hart.— The Geographical Distribution of Life in North America, etc.,” Proc.
Biol. Soc. Wash. vii. pp. 1-64, 1892.

Miter, G. S.—“ List of North American Land Mammals, etc.,” Bull. U.S. Nat. Mus. 1912.
Osporn, H. F.—The Age of Mammals. Macmillan & Co., New York, 1910.

Scuarrr, R. F.—Distribution and Origin of Life in North America. Constable & Co., London,
1911.

Scorr, W. B.—A History of Land Mammals of the Western Hemisphere. Macmillan & Co.,
New York, 1913.

Trovrssart, E. L.—Cat. Mamm., Suppl. 1899-1904.

REPTILIA, BATRACHIA, AND PISCES.
By C. Tate Reaan, M.A.

It has been found convenient to deal with these groups in the reverse order to that
indicated in the heading of the chapter, and to take the Fishes first.

PISCES.

Marine Fishes.—It is well known that the fishes of the Pacific coast of America,
from California to Peru, are quite distinct from those of the tropical Indo-Pacific, and
are related to those of the Antillean district. At one time it was thought that a large
proportion of the species were found on both the Atlantic and Pacific coasts, but the
most recent work tends to show that when pelagic fishes of wide distribution are
eliminated the number of species identical on the two coasts is very small. ‘There
are, however, many closely related species that represent each other on the Atlantic
and Pacific coasts, and the inference is that each pair has evolved from a parent form
that existed when the two oceans were connected, ‘There is evidence that North and
South America were separated by sea during the Eocene, and became one continent
in the Miocene; if this were so, the similarity and degree of dissimilarity between
the tropical shore-fishes of the Atlantic and Pacific coasts would be satisfactorily
explained.

Fresh-water Fishes.—Fishes that are found in lakes and rivers sometimes belong
to marine species that enter fresh water for purposes of breeding or feeding; such
species may form permanent fresh-water colonies or races, and these lead to fresh-water
species of marine genera and fresh-water genera of marine families. These are
unimportant in zoo-geography, but there are many families and even some orders that
are confined to fresh water, and appear to have evolved their genera and species iu
fresh water; their dispersal has depended on hydrographical changes, such as the unioa
of rivers formerly distinct or the capture by one stream of the tributaries of another,
and for most of them the sea appears to be an impassable barrier. Such fishes may be
termed true fresh-water fishes, and they are of the highest importance as indicating

former land-connectious or ancient lines of severance.

The Neotropical Region.—South America has a rich and varied fish-fauna,
surpassing that of any other region for wealth and individuality. As in other parts
of the world, except the Australian Region, the majority of the true fresh-water fishes
belong to the order Ostariophysi*. This order comprises.two well-marked suborders,

* Of, Ann. & Mag. Nat, Hist. (8) viii. 1911, pp. 13-32, 553-577.
BIOL. CENTR.-AMER., Introd. Vol., June 1915. P

106 ORIGIN, ETC., OF THE FAUNA.

Cyprinoidea (Characins, Electric Eels, Carps, Loaches) and Siluroidea (Catfishes).
Of the Cyprinoids the Cypriniformes are absent from South America; the Characi-
formes (Characins) are represented by five families—four endemic and the other, the
most generalized, found in Africa also; whilst the Gymnotiformes (Electric Fels) are
peculiar to the Neotropical Region. The Siluroids include the archaic Diplomystes
of Chile and eight endemic families, of which the most generalized, Pimelodide, is
related to the African and Indian Bagride. The only other Neotropical fishes of much
importance geographically are the Cichlid Perches, also found in Africa, for the
Cyprinodonts are partly marine and the sea has aided in their dispersal.

Origin of N eotropical Fishes.—It has been suggested by various authors that
the ancestors of the characteristic neotropical faunal groups may have reached South
America in late mesozcic or very early tertiary times either from Australia wid
Antarctica, from North America, or from Africa vié an Atlantic continent. Recently *
I have attempted to show that the evidence for the supposed connection with Australia
does not warrant the conclusions that have been reached; so far as the true fresh-water
fishes are concerned, there is only one family common to Australia and South America,
the Osteoglosside, a generalized, ancient, and widely distributed group represented at
the present day by a few remnants. As to North America, it has not, at the present
day, a single family in common with South America, and the fossils show that it has
been just as distinct throughout the tertiary. Cat-fish (Rhineastes), an Osteoglossid
(Dapedoglossus), and a Perch (Priscacara) from the Green River Shales (Lower Eocene)
in Wyoming have been cited as showing neotropical affinities +. In my opinion,
Rhineastes may belong to the cosmopolitan and marine family Ariide, Dapedoglossus
seems to be nearer to the Indo-Australian Scleropages than to the South American
Osteoglossum, and Priscacara is not a Cichlid. All the Cichlid, whether African or
American, have the palate toothless, and the caudal fin formed of 16 principal rays,
14 branched. Priscacara has teeth on the vomer and 17 principal caudal rays,
15 of them branched. After examination of the specimens in the British Museum,
I conclude that Priscacara is a member of the endemic Nearctic family Centrar-
chide, and is closely related to the modern Eupomotis, in which genus the
enlargement and coalescence of the lower pharyngeals may also be seen. If the
ancestors of the present Neotropical fishes came from North America, no trace of
them has yet been discovered.

The hypothesis of a former land-connection between South America and Africa
receives strong support from the Fishes; although no genera of the Characide and
Cichlide are common to the two continents, the close relationship of Brycon and

* Brit. Antarctic ‘Terra Nova’ Exped., Fish. 1914.
T Osborn, ‘ The Age of Mammals,’ p. 136.

REPTILIA, BATRACHIA, AND PISCES, 107

Alestes of the Characide, and of Acara and Paratilapia of the Cichlid, seems
apparent. Further support is derived from the Lepidosirenide, with Lepidosiren in

South America and Protopterus in Africa, and from the relationship of the Pimelodide
and Bagride.

Evolution and Dispersal of Neotropical Fishes.—Since all the genera and
most of the families of true fresh-water fishes of the Neotropical Region are peculiar,
and the majority of the endemic families may be regarded as specialized Characide
or Pimelodide, it may be inferred that the present fish-fauna has evolved in South
America from a few ancestral types, and that it has received no immigrants from other
regions since the earliest of the Tertiary.

It is generally accepted that the Antillean Region was submerged during the Eocene,
and that the invasion of Central America from the south dates back not earlier than
the Miocene. The distribution of the fishes is in harmony with this supposition, for,
whilst most of the Neotropical families have a wide range in South America, only four
of them extend north of the isthmus of Panama. One of these is the Gymnotide,
represented by Gymnotus carapo (Giton fasciatus), found everywhere from Montevideo
to Guatemala; the others are the Pimelodide, Characide, and Cichlide. The
_ Pimelodide are represented mainly by about twenty species of the large and widely
distributed South American genus Lhamdia, which ranges north on the Atlantic side
to southern Vera Cruz, but has not reached the Balsas nor the Mexican Plateau.

A few genera of the Characide have got as far as Rhamdia, and it is only two or
three forms that are scarcely specifically distinct from Tetragonopterus (Astyanaz)
rutilus, found everywhere in South America north of the La Plata, that extend the
range of this family to the Balsas and on the Atlantic slope to the Rio Grande.

In the Characide and Pimelodide invasion of Central America has resulted in the
differentiation of a number of species, but has produced no types that are markedly
distinct from their relatives in South America.

The Cichlide are in many respects peculiar; the Central and South American
species are about equal in number and for the most part are generically or at least
subgenerically distinct, so that it is usually possible to recognize at a glance whether
a species is Central or South American. The South American types are the more
generalized, for the majority have three anal spines and simple conical teeth ; in the
Central American genera the number of anal spines is increased and various
specializations of the dentition occur. The great lakes of Nicaragua have a highly
specialized endemic Cichlid fauna; many species are found in the region between
Panama and southern Vera Cruz, and a few in the Atlantic coast-streams northwards
to the Rio Grande. Only one species, belonging to the genus or subgenus Parapetenia,
is found in the Balsas and a related form in the lowland streams of Sinaloa; a species
of this type also occurs in Cuba. The Central American Cichlide are a difficult group

P2

108 ORIGIN, ETC., OF THE FAUNA.

for the systematist ; the species are closely related and the genera ill-defined. ‘They
give the impression that their number and variety, as compared with the Characide
and the Pimelodide of this area, are not due to an earlier migration, but to a greater
capacity for differentiation and a more rapid evolution.

The Nearctic Region.—North America has a fish-fauna as different as possible
from that of South America. The Ostariophysi are represented by the Catostomide
(Suckers) and by a family of Cat-fishes, the Amiuride, each restricted to this region
except for one or two species in China; in addition, there are a number of Leuciscine
Cyprinide, a group well represented in Eurasia. The Perches belong to the endemic
family Centrarchide and to the Percide. The Cyprinide, Percide, Esocide, and
Umbride are holarctic, but there are several endemic families, Hiodontide, Per-
copside, etc., in addition to those mentioned above, that make this region quite
distinct from the Palearctic.

Origin of Nearctic Fishes.—Numerous fresh-water fishes have been described
from early Tertiary deposits in North America. Lepidosteide and Amiide were
already present in the Upper Cretaceous or Basal Eocene of North America and
Europe. An interesting assemblage is known from the Lower Eocene (Green River
shales) of Wyoming, including Percopside or Aphredoderide (Amphilaga and Erisma-
topterus), Percide (Mioplosus), and Centrarchide (Priscacara), as well as the Osteo-
glossid Dapedoglossus; there are some other fishes that may have been marine as well
as fresh-water, e. g. Xiphotrygon (Trygonide), Notogenus (Chanide), Diplomystus
(Clupeide), and Rhineastes (Ariide). Catostomide are also known from the Eocene
and Amiurus from the Lower Miocene. |

The known history of the Nearctic fishes is in harmony with the supposition that
North America separated from Eurasia at the end of the Cretaceous and developed
several of its endemic types during the Eocene, and that since the Eocene one or
more connections with Eurasia have brought Leuciscine Cyprinide to America, and
have established two or three Catostomids and Amiurids in China. The identity of
some of the more northern species of the two continents shows that a connection
across the Behring Sea must have persisted until a recent date.

Dispersal of the Nearctic Fishes.—In the Mississippi all the characteristic
nearctic types are represented; to the north, west, or south there is a marked
impoverishment. In the Rio Grande, Hiodontide, Percopside, Apbredoderide,
Umbride, and Esocide are absent, and the Percide and Centrarchide are reduced to
a few species only ; there are several Cyprinids and Catostomide. Isolated streams
and lakes of Chihuahua and Durango, the portions of the Yaqui and Mezquital Rivers
east of the Sierra Madre, and the rivers of Tamaulipas and northern Vera Cruz have a
fish-fauna essentially similar to that of the Rio Grande. On the southern part of the
Mexican Plateau, the Rio Grande de Santiago or Lerma above the falls, with the

REPTILIA, BATRACHIA, AND PISCES. 109

isolated lakes of the states of Michoacan and Mexico, constitute an important system
with a very characteristic fish-fauna formed almost entirely of endemic species. Only
three families of Nearctic fishes are present; the Catostomide are represented by a
single species of Mowostoma and the Amiuride by an Amiurus, whilst there are
a dozen Cyprinide, nearly all belonging to endemic genera. The majority of the
fishes belong to two groups, each with about a score of species, and each belonging to
a family that includes a number of marine fishes that enter fresh water. ‘These two
groups are characteristic of the Lerma System and barely overstep its limits; one is
the somewhat heterogeneous Atherinid genus Chirostoma, the other is the Cyprinodont
subfamily Characodontin, which includes five well-marked genera, and differs from
the Funduline in that its species are viviparous.

The Lerma System has evidently long been isolated, and has been a centre of
evolution. The chain of volcanoes that border the Mexican plateau to the south have
barred invasion on that side, and from the north only a few nearctic types have
reached the Lerma. The main elements of its fish-fauna, Chirostoma and the Chara-
codontine, have in all probability evolved each from a single ancestral type that
entered from the sea, from which it is now separated by inaccessible falls.

The Rio Balsas lies to the south of the Mexican plateau, and is a large river that
flows into the Pacific; it has not been thoroughly investigated, but appears to have
a comparatively poor fish-fauna. In addition toa Lerma type (Goodea) that has got
into the headwaters of its northern tributaries, there is a Cat-fish (Amiwrus) and a
Cyprinoid (Notropis) ; two neotropical species balance these. The high mountains
that bound the Balsas System have evidently rendered immigration a difficult feat.

It is in the lowlands of the Atlantic slope that conditions have been more favourable
to migration. Lepidosteus, absent from the Mexican plateau and from the Balsas, has
reached Panama, and a species of Ictrobus (Catostomidee) and an Amiurus occur in
the Usumacinta, balancing the Cichlid and Characid species of the Rio Grande.

Dispersal of Neotropical and Nearctic Fishes compared.—tIt has been
shown above that comparatively few elements of the rich neotropical fish-fauna extend
north of the Isthmus of Panama, and that of these only the Cichlide exhibit much
diversity and specialization. In Central America and in rivers of the Atlantic slope
of Mexico south of Vera Cruz the fish-fauna is almost exclusively neotropical, but
only two neotropical types have reached the Balsas, none have surmounted the volcanic
chain that borders the Mexican plateau, and only half a dozen have penetrated north
of Vera Cruz in the Atlantic coast-streams of northern Mexico. In these streams
there are many nearctic species, but in Atlantic rivers south of Vera Cruz only three.
The Rio Grande exhibits a paucity of nearctic types as compared with the Mississippi,
and this is still more marked in the Lerma System, which possesses a peculiar endemic
fish-fauna. In the Balsas, equally isolated from the north and the south, nearctic and
neotropical species balance, but there are only two or three of each.

110 ORIGIN, ETC., OF THE FAUNA.

It is probable that the distinctness of the North and South American faunas 1s
more marked, and that where they meet there is less overlapping in the case of the
fresh-water fishes than in any other group of animals.

Fresh-water Fishes——General Conclusions.—The South American fresh-water
fish-fauna is as distinct as the Mammalian fauna would have been if a large part of
the endemic fauna had not died out in late Tertiary times, and been replaced by the
invasion of types that had evolved elsewhere.

Turning to the other parts of the world, we find that the distinctness of the
Australian Region, marked in the Mammalia, 1s emphasized by the Fishes. ‘The
effectiveness of the sea asa barrier is strikingly illustrated by a comparison between
Borneo, with its hundreds of species of Cyprinoids, Siluroids, Anabantoids, etc., and
Celebes, without a single indigenous true fresh-water fish. The other four regions are
well characterized, although less distinct, for the Indian has certain relationships with
the Ethiopian and Palearctic, and the last with the Nearctic ; there has been a certain
interchange, but in each case the endemic groups preponderate.

The general impression derived from a study of the fresh-water fishes of the world is
that many of the families were in existence at the beginning of the Tertiary, that for the
most part they have evolved in the areas they now occupy, have dispersed slowly, and
have never had a distribution much wider than at present. For Mammals, it may be gene-
rally stated that they have evolved rapidly and spread rapidly, and have found it easier
to reach a country than to live in it. For fresh-water fishes migration has been much
more difficult, survival relatively easy ; hence their great importance in zoogeograpby.

BATRACHIA.

Batrachians.—These resemble fresh-water fishes in their inability to swim across
the sea, and their eggs are no more likely to be transported over the sea than those of
fresh-water fishes. But it is probable that accidental transmission has played a part in
the dispersal of some arboreal and terrestrial frogs ; at any rate, this seems the most
reasonable explanation of the distribution of certain genera and species in the Indo-
Australian Archipelago. Even the most aquatic types can travel overland from
one stream to another, and consequently are able to migrate rapidly when conditions
are favourable.

Nearctic and Neotropical Batrachians.—The Batrachian faune of these two
regions are less distinct than the fish-faune, but are, nevertheless, very different.
Urodeles are scarce in South America, Ceecilians are absent from North America; the
majority of the Neotropical Frogs are Cystignathide and Hylide, families but sparingly
represented in the Nearctic Region.

The mountains that fringe the Mexican Plateau form the boundary between the

REPTILIA, BATRACHIA, AND PISCES. lil

the Nearctic and Neotropical regions; only a small proportion of the tropical genera
have gained a footing on the plateau, and but a few undoubted northerners have
spread far to the south beyond it.

The Urodela are almost exclusively holarctic and aquatic; except the monotypic
Thorius, known only from the mountains of Orizaba and Oaxaca, but two genera, Ambly-
stoma and Spelerpes, extend southwards into Mexico and Central America. Each of
these genera has several species in the United States, and whilst Amdlystoma has one
in the mountains of Siam, Spelerpes includes one from Sardinia and Italy. These
isolated species suggest that both genera had formerly a wider and a more northerly
range, and that only one Old World species of each managed to survive the glacial
epoch by migrating southwards. In America Amblystoma extends on to the Mexican
plateau and the mountains that fringe it, but Spelerpes includes about seven species
from the United States, fifteen from the mountains of southern Mexico and Central
America, three from the Andes of Peru and Colombia, and one from Haiti. It seems
evident that this genus began to spread southward as soon as the elevation of Central
America enabled it to find a congenial temperature at high altitudes.

The Cecilians are vermiform, apodal, burrowing animals that inhabit the Neo-
tropical, Ethiopian, and Indian regions. There are a few Central American species,
and one of these, Dermophis mexicanus, ranges north to southern Vera Cruz. Nothing
is known as to the past history of this group, and they may or may not be ancient

inhabitants of South America.

The Anura, or Frogs and Toads, may be aquatic, terrestrial, arboreal, or fossorial.
Some genera and species have an extremely wide range, and, although it is evident
that the sea forms an impassable barrier for the majority, it seems probable that
accidental transmission has played a part in the dispersal of genera such as Rana
and [Hyla.

The two main divisions of the Anura—Firmisternia and Arcifera—may or may
not be natural, but the further classification into families is unsatisfactory; the
Cystignathide, for example, are heterogeneous, as the Australian genera are very
distinct from those inhabiting South America, the latter appearing to form a natural
group. ‘The Bufonide also may be an artificial assemblage.

To the Firmisternia belong the Ranide and Engystomatide, abundant in South
America, Africa, India, and ranging eastwards to New Guinea and beyond. There
are several endemic South American genera, and two or three with Central American
species also. Hngystoma extends from Paraguay to Texas and Florida. Hypopachus
has one species ranging from Brazil to Costa Rica, another from Ecuador to
Guatemala, and a third from Western Mexico to Texas. The only other genus of
the group found in North America is the nearly cosmopolitan Rana, which extends

112 ORIGIN, ETC., OF THE FAUNA.

southward to the Andean region of South America, and is doubtless a recent
immigrant. Gadow suggests that on reaching South America it gave rise to the
endemic neotropical genera referred to the Ranide, but it is possible that these are
derived from an ancient neotropical stock, for at least one recent genus of the Ranide
dates back to the Eocene, and the family itself may have a greater antiquity.

Of the Arcifera, the American Cystignathide are almost exclusively neotropical ;
there are several endemic South American genera, and of those that range northwards
into Central America only Syrrhophus, Hylodes, Leptodactylus, and Borborocetes are
represented on the Mexican plateau by a few species ; none range further north.

The Hylide are mainly neotropical and arboreal, and except for one or two species
of Hyla are absent from the Mexican plateau. Only Hyla ranges outside this region
through North America to temperate Eurasia and southwards through China to
Burma, then comes a gap, for the Australian Region is rich in species, only one of
which crosses Wallace’s Line into the Sunda Islands. The presence of Hyla arborea
in Madeira and the Canaries, the range of H. dolichopsis from Sumatra to the
Solomons, suggest that these tree-frogs may be carried across the sea, and that their
wide distribution may be partly due to this; their absence from Africa and India is
remarkable, but there are other equally curious cases of discontinuous distribution
in genera of no great antiquity, and it is unnecessary to infer some special connection
between Australia and South America to explain this one; nor does the distribution
of the Cystignathide support this view, for the Australian members of the family are
by no means closely related to the American ones, and their common ancestor must
date back well into mesozoic times.

Of the American genera referred to the Bufonide two are strictly neotropical,
Engystomops ranging from South America to Tehuantepec, and the monotypic
Rhinophryne occurring on the Atlantic slope from Vera Cruz to Guatemala. The
third genus, Bufo, is cosmopolitan except for the Australian Region; there are a
number of species in Central America and southern Mexico, whilst others on the
Mexican Plateau range into the southern United States. Except for two species in
Celebes, Bufo is absent from the Australian Region, although there are many species
in the Malay Peninsula and Archipelago; this is exactly the reverse of what occurs in
Hyla, and Rana differs from both, as it is well represented in both Indo-Malayasia
and Austro-Malayasia eastwards to the Caroline Islands.

Scaphiopus, the American representative of the family Pelobatide, is related to the
European Pelobates; there are about eight species from the United States and the
Mexican Plateau ; these are diggers in the sand, and do not range southwards beyond
the mountains of Southern Mexico.

Dispersal of Batrachians,—The Anura and Urodela are both known to date
back to the Jurassic, and this, as well as the presence in New Zealand of a Toad

REPTILIA, BATRACHIA, AND PISCES. 113

(Liopelma), indicates that in all probability some of the Batrachian families are more
ancient than those of fresh-water fishes. There seems no reason to doubt that they
can and have spread rapidly, but probably their main distribution was accomplished
before the Tertiary. The Urodela have in all probability always been northerners,
and have only recently penetrated into South America. ‘The Cecilians are tropical,
but their absence from the Australian Region rather tells against the idea that they
are a very ancient group. Many families of the Anura are widely distributed

and probably ancient; the Cystignathide and Hylidae seem to be old neotropical
families.

REPTILIA.

Reptiles.—Leaving out the aquatic members of this group, some Reptiles can and
do swim across the sea; others that are arboreal or that burrow into or hide in trunks
of trees are peculiarly liable to accidental transmission over the ocean. Nearly all
can travel quickly overland, so that in all probability the dispersal of many groups
has been accomplished rapidly. The difference between the Nearctic and Neotropical
regions is decidedly less marked than in the Batrachians, but the boundary is
the same.

Of the groups of Reptiles represented in Central America the Crocodiles and
Tortoises are ancient and seem to have been cosmopolitan at the beginning of the
Tertiary ; a study of their present distribution leads to no very important results.
The Lizards and Snakes are the dominant orders of the present day, but their past
history is almost unknown; in all probability the principal families date back to early
FKocene times, and as they developed during the Tertiary they may have used the same
routes of migration as the Mammalia.

Dr. Gadow’s papers on the Reptiles and Batrachians of Mexico and Central
America have been referred to above (p. 59) and are of great interest, especially
when they deal with the effects of altitude, climate, terrain, etc., on the distribution ;
his deductions as to the origin and migrations of the different groups are necessarily
based mainly on their present distribution, and are to that extent unsatisfactory. The
Mammals, whose evolution during the Tertiary, extended migrations, survivals, and
extinctions are very thoroughly known for several orders, teach that it is very unsafe
to deduce centres of origin and dispersal from present distribution alone. —

Lizards.—There appear to be no families of Lizards restricted to South America,
but most of the 35 genera of one family, the Teiide, are peculiar to that continent ;
half a dozen South American genera range into Central America, two reaching

southern Mexico, and one—Cnenudophorus—the United States. This genus includes
, species on the Mexican plateau, as well as in the lowland forests. ‘The conclusion
that this is an old neotropical family seems not unreasonable, but may be incorrect ;
the neotropical Monkeys furnish a parallel. | |

BIOL. CENTR.-AMER., Introd. Vol., June 1915. Q

114 ORIGIN, ETC., OF THE FAUNA.

The family Amphisbenide comprises burrowing Lizards, with limbs reduced or
absent; the majority of the genera are neotropical or African, but the most
generalized genus, and the only one with fore limbs, Chirotes, is found in California
and Mexico, and the least specialized of the rest, Blanus, inhabits the countries round
the Mediterranean. Moreover, Rhineura of Florida is known to date back to the
Oligocene of Dakota. Here an American distribution similar to that of the Teiide
is coupled with indications that the group may have been originally a northern one
that migrated southwards into Africa and South America.

The Scincide, widely distributed in the Old World, have in all probability reuchnd
America quite recently, as the three genera found in America are also Asiatic.
Eumeces has several North American species, and extends southward over the plateau to
southern Mexico. Lygosoma laterale of the southern United States ranges southward
to southern Mexico, and in Guatemala is replaced by the closely related L. assatum.
L. laterale is almost identical with Z. reevesi of China and Burma, and it may be that
in the warmer climate of Pliocene times the ancestral type ranged northward to the
land connecting Asia with America, The third American genus, Mabuwia, is found
also in Africa and southern Asia; there is an Antillean species, one from Costa Rica,
and four from South America, one of these extending northward to Yucatan and
southern Vera Cruz. This neotropical distribution of a genus that in all probability
came through North America parallels that of Zapirus.

The Iguanide are American, except for two genera in Madagascar and one in the
Friendly and Fiji Islands. They are also known from the Eocene of Europe, and are
evidently an ancient group of former wide distribution; the presence of two peculiar
genera in the Galapagos suggests their antiquity in South America. The South
American genera are numerous; the majority are restricted to South America, but
some are also Antillean, and some extend through Central America as far as southern
Vera Cruz; there are also two or three genera peculiar to Central America. ‘There
are about ten genera in the south-western United States, some with species on the
Mexican Plateau and the western Sierra Madre. One species of Uta (U. bicarinata)
is found on the plateau and also on the west coast from Presidio to Tehuantepec.
Sceloporus and Phrynosoma are important genera that range from the southern United
States to Central America, and include species on the plateau as well as in the
lowlands.

The Geckonide are found in all tropical countries; in America they are almost
entirely neotropical, and are not found on the Mexican plateau. ‘The Geckonide are
peculiarly liable to accidental dispersal, and this is well illustrated by the distribution
of Gehyra mutilata ; it is found on the islands of the Indian Ocean and the Malay
Peninsula and Archipelago eastward to New Guinea, but it is also known from
Western Mexico. The small family Eublepharide, specialized Geckos, includes three
species from Africa and India, distinguished by their stout form and by their enlarged

REPTILIA, BATRACHIA, AND PISCES. 115

chin-shields from the four American species ; two of these, from Texas and California
and from the West Coast of Mexico, differ in their scaling from the other more
tropical species, one of which inhabits Panama and the other ranges from Costa Rica
to Vera Cruz.

The other families of Lizards found in America teach us nothing more than those
dealt with above. There are several families common to the nearctic and neotropical
regions, and most of these are found in other parts of the world also. South America
is much richer in genera and species than North America; most of the Lizards of
Central America north to the outer slopes of mountains that border the Mexican
plateau belong to or are related to South American genera, and most of those found
on the plateau are generically distinct from them, but in some cases (e. g. Sceloporus,
Phrynosoma) only specifically so.

There is not, on the whole, a very fundamental difference between the Lizard faune
of North and South America; considering the differences in climate, and other
features, we should expect most of the genera to be distinct, and in southern Mexico
the transition from an elevated, dry, and almost treeless plateau bordered by mountains
to a lowland district with tropical rainfall and vegetation accentuates this distinctness.
But there are few differences that can be assigned definitely to the former isolation of
the two continents, as practically nothing is known of the Tertiary history of the
group. We know that some Mammalian genera, now tropical, formerly ranged
further north and migrated from Asia to North America and thence to South America
in late Tertiary times. Doubtless some Lizards, such as Mabuia, had a similar
history ; but whether the Teiide, for example, were formerly nearctic, or whether
they are for the first time spreading northwards, is a question difficult to answer.

Snakes.—The burrowing snakes of the genus Typhlops have a wide distribution in
the tropics; there are a few species in Mexico and Central America, and one of these
is ZT. brahminus, collected by Dr. Gadow in Michoacan. Previously this species was
known to range from Africa to China and on islands eastward to the Moluccas;
probably it will be found on some of the islands of the Pacific. ‘This is an interesting
parallel to the Gecko Gehyra mutilata ; in both cases Indian species have reached the
West Coast of Mexico, and appear to have established themselves. The occurrence of
Typhlops on oceanic islands indicates that the wide range of this genus is not due to
its antiquity, but to susceptibility to accidental dispersal.

Another genus of burrowing suakes, Glauconia, is found in America, Africa, and
south-western Asia. (G. albifrons is tropical, ranging from Argentina to southern
Mexico, but other species are found on the Mexican plateau and in the southern
United States.

The Boide show some points of interest. The Pythonine include a score of species
from Africa, India, and Australia and a single American species, Loxocenus bicolor,

Q 2

116 ORIGIN, ETC., OF THE FAUNA.

from Guatemala and southern Mexico. The Boine are widely distributed in the
Old World, but the majority are American, and the occurrence in Madagascar of
species of the ‘Tropical American genera Corallus and Boa is curious; it is evident
that these genera had formerly a much wider range.

Many other genera of snakes have a wide distribution; thus the Crotalide have
several genera common to Asia and America, and, although they extend southwards
to the La Plata, there are no peculiar Neotropical genera. One species of Crotalus
ranges from the United States to Argentina.

Without going into further details, it may be said that, as compared with the Lizards,
a larger proportion of the Central American genera are found also on the Mexican
plateau and in the southern United States, although there are, of course, a number of
neotropical genera that range northwards only to southern Mexico.

The Amblycephalide are the only family of snakes that are strictly neotropical in
America; there are three South American genera, one of these, Leptognathus, having
several Central American species, the northernmost in Tehuantepec. The other two
genera occur in South-Eastern Asia, and it is probable that the family was formerly
a northern one. Indeed, it is not unlikely that the whole neotropical Ophidian fauna
has been derived from immigrants from the north that did not begin to reach South
America until the Miocene connection was established. ‘The distribution of the vast
family Colubride seems to favour this view.

REPTILIA, BATRACHIA, anp PISCES.

Summary.—The mountains that fringe the Mexican Plateau form the boundary
between the Neotropical and Nearctic Regions. True fresh-water fishes can neither
cross seas nor travel overland from one river to another ; consequently their dispersal
has been slow. ‘The nearctic and neotropical fishes are quite different, and belong to
different families; the two faune scarcely overlap; in the Lerma System, on the
southern part of the Mexican plateau, there are no neotropical fishes, the nearctic
fishes are mostly generically distinct from those of the Rio Grande, and two endemic
groups that are neither nearctic nor neotropical are the most important elements.

For most Batrachians the sea is an impassable barrier, but they can migrate over-
land and their dispersal may have been rapid. The nearctic and neotropical
Batrachians are mostly distinct, but overlap to a considerable extent; a few genera
(Bufo, Hyla) range throughout both regions. The rapidity of dispersal as compared
with the fishes is exemplified by the holarctic group Urodela; one genus, Spelerpes,
has penetrated far into South America, whereas only one nearctic fish, the ancient
Lepidosteus, has reached Panama. The spreading of Amblystoma tigrinum. of the
United States southwards to the border of the Mexican plateau is in striking contrast to
the peculiarity of the fish-fauna of the southern half of the plateau, where no species

REPTILIA, BATRACHIA, AND PISCES. 117

of fish occurs that is found in the United States. Crocodiles and Tortoises, like many
families of Anura, are ancient and widely distributed. Snakes and Lizards are more
modern, and their dispersal seems to have been recent and rapid; in these groups
there is no marked difference between the nearctic and neotropical regions; no
orders or families that show much regard for the boundary; the differences are
for the most part generic or specific, and are mainly due to the sudden change
from the dry plateau of the southern United States and Mexico to the tropical
forests of Central America. .
Of the groups dealt with in this chapter, only the Fishes give clear indications as to
the past, and from them we infer a Cretaceous connection between South America and
Africa that may have persisted into the Eocene, and was succeeded by a long isolation
of South America and in the Miocene by its connection with North America, which
has been since the Eocene until quite recently more or less continuously connected
with Asia across the Bering Sea. |

[ 118 |

ARACHNIDA *.
By R. I. Pococs, F.R.S.

INTRODUCTION.

Discussions on the geographical distribution of the Arachnida, especially of
Scorpions, commonly open with a reference to the great antiquity of the group, and
to tle possibilities thus afforded it of achieving cosmopolitan dispersal. Dr. Hans
Gadow, for instance +, dismisses the Scorpions in the following paragraph :—“ ‘This
group is a good illustration of the effect of great antiquity. Scorpions already existed
in the Silurian, and even some existing species date back to the Coal Measures!
‘They have had every chance of spreading widely. A species of Tityus is preserved in
Miocene amber of the Baltic, this genus is now restricted to southern South America.
The group is cosmopolitan, limited only by cold, yet it is absent from New Zealand.
They show scarcely any generic affinity between the Old World and the New, nor
between South America and Australia. They have had sufficient time to develop
along lines aloof from each other in these great land complexes.”’

Much of this is untrue both in substance and in fact. None of the many Carboni-
ferous genera can be referred with certainty even to existing families; and the record
of a species of “ Zttyus” from the European Miocene can only be regarded as evidence
of the existence of the Buthide in the Baltic area in mid-Tertiary times; and, as
regards the distribution of existing forms, if the views above expressed be accepted,
there is nothing more to be said upon the subject. They are, however, inadmissible
since they leave wholly unexplained the fact, singular though it be, that the present
distribution of Scorpions does not attest the great antiquity of this order. If they
were not known to be of Carboniferous age they might be judged, from the analogy
supplied by Mammals, to date from late Mesozoic and early Tertiary times, for, as I
have elsewhere pointed out {, if the surface of the world be regionally divided in
accordance with the distribution of Scorpions, the resulting map will agree tolerably
closely with the map based upon the distribution of Mammals, due allowance being
made for the absence of Scorpions at the present time from all countries to the north,
roughly speaking, of the 45th parallel of north latitude.

* The Opiliones and the Acari have been omitted from this essay, the former because Mr. F. O. P. Cam-
bridge was not sufficiently acquainted with them to make his results altogether reliable, and the latter
because of the imperfection of our knowledge of the Acarine fauna of other parts of the world.

+ ‘The Wanderings of Animals,’ 1913.

~ ‘ Natural Science,’ iv. pp. 853-364, 1894, and 1899, pp. 213-231.

ARACHNIDA. 119

A striking Mammalian feature in the distribution of Scorpions is their absence from
New Zealand. Another is to be found in the character of the Australian forms, some
of which are quite peculiar, while others show affinity with genera and species from
southern Asia, like the Australian Rodent Mammals, and one, like Thylacinus, belongs
to a South American family. Again, some of the South Asiatic genera do not pass
Wallace’s Line, and the small number of Mascarene forms are related to, but distinct
from, those of tropical Africa and Asia, like the Mascarene Civets and Lemurs.
Similarly, the tropical African and Asiatic species, although related, belong mostly to
distinct genera.

What is true of the Scorpions is true also, broadly speaking, of the two suborders of
Pedipalpi, the Urotricha and the Amblypygi, which, like the Scorpions, existed in
Carboniferous times in Europe and North America. The absence of both from New
Zealand and Australia, and of the Urotricha from Africa also, are suggestive rather of
comparatively recent origin than of high antiquity.

The same cannot, however, be said of a great many of the Aranew (Spiders). The
most ancient existing type, Liphistius, apparently related beyond doubt to Carboni-
ferous genera, is restricted to Indo-Malaysia, whereas many genera, like Lycosa,
Aranea, Tetragnatha, and others, not known to be ancient forms, are practically
cosmopolitan in distribution. Moreover, some groups of spiders—perhaps Mesozoic or
even Czenozoic, but not, so far as records tell us, Paleeozoic—attest by their restriction
to the southern Continents the former existence of ‘‘ Antarctica” much more forcibly
than the orders of Arachnida known to have been in existence in the Carboniferous
Period. Another instance of the restricted distribution of a Carboniferous order is
supplied by the Rianulei, which are found now only in tropical West Africa and Brazil.

Thus some of the Carboniferous Arachnida, the Scorpions, exist in Europe, Asia,
Africa, America, and Australia; others, the Amblypygi, in Asia, Africa, and America ;
others, the Urotricha, in Asia and America; others, the Rianulei, in Africa and South
America; the Mesothelid Spiders (Liphisticus) only occur in the East Indies, while
other spiders, assumed on morphological grounds to be of later date, may be cosmo-
politan, and such spiders, when young, are known by their method of floating on
webs to be able to cross arms of the sea in the direction of prevalent winds *.

From these facts it seems clear that the present distribution of the Arachnida
depends, not upon the duration of their existence, but upon their means of dispersal
and power of adaptation to varied conditions. Hence the attempt to explain away
the facts of their distribution by an appeal to their antiquity is barren of results, and,
since the paleontological history of the Arachnida throws no useful light upon the
matter in hand, it has been ignored in the following discussion.

* This means of dispersal is analogous to the flight of birds, whose powers to cross tracts of ocean cannot

be ignored in discussions of geographical distribution.

120 ORIGIN, ETC., OF THE FAUNA.

Order SCORPIONES *.

The genera of this order recorded from Central America may be classified, with
their distribution, as follows :—

Family ScorPionipz.

Subfamily Ischnurine.
Genus Opisthacanthus. Panama, Colombia, W. Indies.

Subfamily Diplocentrine.
Genus Diplocentrus. Texas, Mexico, W. Indies, Brazil.

Family Vasovip2.

Subfamily Vejovine.
Genus Hadrurus. S. States of N. America ; Mexico, ? Guatemala.
Genus Vejovis. 8S. States of N. America; Mexico.
(Genus Uroctonus. California; ? Guatemala.)
(Genus Anuroctonus. California, Utah, etc. ; ? Guatemala.)

Subfamily Megacormine.
Genus Megacormus. Mexico.
Genus Plesiochactas. Mexico.

Subfamily Chactinz.
Genus Broteochactas. Panama, Guiana.

Family Buruipz.

Genus Centruroides. S. States of N. America, Central America, Antilles, S. America.
(Genus Rhopalurus. Cuba, Haiti, Brazil, ? Mexico.)
Genus Jityus. Mexico, Costa Rica, Panama, W. Indies, S. America.
(Genus Isometrus. Commercially imported, if present.)

With the substitution of the older name Vejovide for Iuride and the introduction
of the Megacormine, containing genera unknown to me at the time, this is the
classification proposed by myself in 1893 (Ann. Mag. Nat. Hist. (6) xii. pp. 305-311)
- and adopted in my paper on ‘“‘ The Geographical Distribution of Scorpions” in 1894
(Nat. Science, v. pp. 353-364). In his contribution to the same subject published in
1905 (Zool. Jahrb. Syst. xxii. pp. 321-364), Kraepelin suggested certain modifications,
erecting the Diplocentrine, Vejovine, and Chactine to full family rank, including the
Megacormine as a subfamily of the Chactide, and making Centruroides, with
Rhopalurus, and Tityus, the types of separate subfamilies of the Buthide. These
amplifications, whether desirable or not, are of no importance in the present
connection.

Some of the genera above enumerated have been placed in brackets because

e

* Although it is customary to designate the primary divisions of the Arachnida as “ Orders,” it must be
remembered that the morphological differences between the groups 0 of that rank are greater than those which
distinguish the ‘‘ Classes” of the Vertebrata. : :

ARACHNIDA. 121

the evidence for their occurrence in Central America is doubtful and requires
confirmation *.

DISTRIBUTION OF THE FAMILIES AND SUBFAMILIES OF
CENTRAL AMERICAN SCORPIONS.

No genera of Scorpions are common to the Old and New Worlds, except Jsometrus,
one species of which has been commercially carried everywhere from the East Indies.
None of the families mentioned above, however, are peculiar to America. The Old
World appears to be the headquarters of the Scorpionide, since this family is
represented there by many genera distributed over Africa, South Asia, and Australia
and referable to four subfamilies, only two of which, the Ischnurine and Diplocentrine,
occur also in America.

The Veejovide, absent from Africa and Australia, range from the Mediterranean
area eastwards through North India into Further India, and reappear in North,
Central, and South America, but are absent from the Antilles. Unless the Mediter-
ranean genus Jurus and the Indian genus Scorpiops be included, as has been done,
in the Viejovine, the American subfamilies are peculiar to the New World. But, in
any case, the distinctions between the subfamilies of this family are finely drawn.
Excluding the two Old World genera above mentioned, the Vejovine contain the four
American forms enumerated above and two from South America. No other genera of
the Megacormine are known. The Chactine, however, are represented by several
genera in the northern countries of South America.

The Buthide are mainly an Old World group. The family contains a large number
cf genera and species in Europe, Africa, Madagascar, South Asia, and Australia.
Apart from the aberrant South American genus Ananteris, the only known American
genera are the three indigenous forms enumerated above. These do not differ more
from each other, or from some of the Old World genera, than the latter differ from
each other. ‘There is no evidence, indeed, that Zityus is not closely allied to the

Oriental genus Jsometrus.

* The genera Uroctonus and Anuroctonus, for example, were included in my report on the testimony of
Thorell, who recorded Uroctonus mordax and Anuroctonus pheodactylus, two species originally described from
California, from Guatemala. The specimens he had for examination, however, were sent to him with an‘
example of Hadrurus hirsutus, also a Californian species, by Dr. Gustav Eisen, who was then living at
San Francisco, and my opinion that they were wrongly localised is based upon the improbability of identical
species inhabiting the two countries in question. The Buthid genus Rhopalurus was included on the evidence
of old specimens of Rhopalurus junceus, ticketed “* Mexico ” in the British Muscum, the species in question
having its home in Haiti. Finally, although Isometrus no doubt exists in the seaport towns of Central
America, it has not yet been actually recorded, and as a commercially introduced form is wholly without

interest.

BIOL, CENTR.-AMER., Introd. Vol., June 1915.

122 ORIGIN, ETC., OF THE FAUNA.

GENERAL FEATURES OF THE SCORPION FauNA OF CENTRAL AMERICA.

The nine genera of Scorpions certainly ascertained to be Central American are
faunistically assignable to a northern category, a southern category, and a category
which is both northern and southern.

To the southern category belong Opisthacanthus, occurring in Panama and Colombia ;
Broteochactas, also found in Panama and the northern countries of South America
(Guiana, etc.); and Zityus, which ranges all over South America from the Argentine
northwards and is found in Panama, Costa Rica, and Mexico. ‘The Mexican species,
however, is very little known. It is not closely allied to the species from Costa Rica
and Panama, which are identical with, or very nearly related to, species from the
Amazons, Demerara, etc. Similarly, the species of Opisthacanthus and Broteochactas
occurring in Panama are the same as species found in South America.

To the northern category belong Hadrurus and Vajovis, which extend from the
Southern States of North America into Mexico, Vajovis being one of the dominant
Scorpions in the latter country. It is interesting to note the wide geographical
severance between these two and their nearest allies, Caraboctonus and Hadruroides,
which inhabit Ecuador, Bolivia, and Chile. Also to this northern category must be
referred the two genera of Megacormine, Megacormus and Plesiochactas, which are
found in Mexico (Vera Cruz, Cordova), the latter extending to Guatemala. Their
nearest allies are the Chactine of tropical South America.

To the category which is both northern and southern in distribution belong the
genera Diplocentrus and Centruroides. ‘The former ranges from Texas into Mexico,
but not, so far as is known, to the south of the latter country in Ceutral America.
Nevertheless, it is found in the Greater and Lesser Antilles and in the northern part
of South America (Brazil). Centruroides, on the other hand, occurs in the southern
states of North America, throughout Central America, in South America as far down
as Chile along the western side, in Brazil, and in the Greater and Lesser Antilles,

Thus of the Central American genera only four occur in the Antilles, Tityus,
Centruroides, Diplocentrus, and Opisthacanthus.

Iwo of the above-mentioned genera, Opisthacanthus and Diplocentrus, are of
outstanding interest from the geographical point of view, because of the singular
distribution of the subfamilies to which they belong.

Opisthacanthus is unmistakably related to the tropical and South African genus
Opisthocentrus. The generic distinction between the two is not, indeed, always
admitted. ‘Tropical Africa is also the home of several more genera of the subfamily
Ischnurine, while others occur in Madagascar and South Asia as far east as
Australasia. The kinship between the African and the solitary monotypical tropical
American genus suggests that South America received this element of its fauna by
means of a direct land-connection with Africa.

ARACHNIDA. 123

The distribution of the Diplocentrine is still more remarkable. Four genera belong
to this subfamily, two being American and two Arabian. In addition to Diplocentrus *,
the range of which has already been stated, America contains the well-marked genus
Oiclus, known hitherto only from Antigua in the Lesser Antilles. The Arabian genera
are Nebo recorded from Syria and Arabia and Heteronebo discovered in the island
of Abd-el-Kuri by Sokotra. This singular discontinuity in distribution is closely
paralleled by that of the Centipede Scolopendra (see p. 134). To explain it one may
tentatively suggest a transatlantic connection between the Mediterranean area of
Europe and America by way of the Antilles, an hypothesis supported by the survival
of Scolopendra in the Canary Islands. Assuming this to be the explanation, it is
probable that America received its Diplocentrine Scorpions from the Mediterranean
area, and not vice versd, because the Old World, and not the New World, is at the
present time the home of the Scorpionide, to which the Diplocentrine belong.

Very little can be said about the origin of the remaining Central American genera.
The absence of the Vejovide from tropical Africa eliminates the latter country as a
possible source for this family. Its mainly. extratropical range in the Old World,
coupled with its existence at the present time in North America, suggests its former
wide distribution in the Northern Hemisphere; and possibly the break in the
distribution of the Vejovine in America, above alluded to, points to the southward
migration of the family from the western parts of North America to the western parts
of South America before the elevation of the isthmus of Panama. In this connection
the absence of the family from the Antilles is significant.

The dominance of the Buthide in the Old World as compared with the New
World suggests the Old World as the original home of the family; but whether the
ancestors of the American genera entered America by a transatlantic bridge from
Africa or Europe or by the known eastern or western northern routes under tropical
or warm temperate conditions, it is impossible to say.

REGIONAL AND SUBREGIONAL DISTRIBUTION OF AMERICAN SCORPIONS.

Making allowances for incomplete knowledge, which renders accurate mapping of
boundaries impossible, it seems that America may be divided by its Scorpions into the
same geographical areas as those indicated by the Mammalia. ‘The northern parts
of the continent to the north, roughly speaking, of the 40th parallel agree with the
corresponding areas of the Old World in being without Scorpions. This negative
feature defines the Holarctic Region. Sharply distinguished from this by the
presence of Scorpions are the Southern States of the Union ; and the extension of two
of the genera (Hadrurus, Vejovis) into Mexico, but no farther, affiliates the greater part

* Including Didymocentrus, Kraepelin.

R2

124 ORIGIN, ETC., OF THE FAUNA.

of that country with the southern United States to constitute the Sonoran Region,
which corresponds with the Mediterranean Region of the Old World. Two more
genera, Uroctonus and Anuroctonus, are peculiar to the Sonoran, but have not been
recorded from Mexico. Probably also Plesiochactas and Megacormus from Mexico and
Guatemala belong to it. The other Sonoran genera, Diplocentrus and Ceutruroides,
extend far to the south, and link the Sonoran Region with the Antilles and not
South America.

South America, the Neotropical Region, is characterised by the family Bothriuride,
ranging from Peru, Chile, and Brazil to Patagonia; by a large number of genera and
species belonging to the Chactine, mainly restricted to the northern parts of the
continent, one genus Broteochactas spreading into Panama; by the Ananterine, a
peculiar subfamily of Buthide ; by the Buthoid genus Tityus, which spreads through
Panama and Costa Rica to Mexico; by the peculiar genus of Ischnurine, Opisthacanthus,
which occurs in Colombia and Panama; and by two peculiar genera of Vajovine,
Caraboctonus and Hadruroides, ranging from Chile to Ecuador. Thus the southern
portion of Central America contains Scorpions generically, in some cases specifically,
identical with those of the northern countries of South America. Ifit be separated
as a subregion from South America, an immaterial point, it may be defined by the
absence of a large number of South American genera.

The affinities of the Antilles are certainly with Neotropical rather than with the
Sonoran Region. ‘Two Sonoran genera, Diplocentrus and Centrurotdes, it is true, occur
both in the Greater and Lesser Antilles, but both are found in South America as well.
The genus Rhopalurus, allied to Centrurotdes, and occurring in Haiti and Cuba, is
also found in Venezuela, Brazil, and Colombia; Opisthacanthus, inhabiting Colombia
and Panama, has been recorded from Haiti; and the essentially Neotropical genus
Tityus exists both in the Greater and Lesser Antilles. One genus, namely Ocelus, is
peculiar to the Lesser Antilles*. Judged from their Scorpion fauna, therefore, the
Antilles may be regarded as a subregion of the Neotropical Region, characterised
mainly by the absence of some conspicuous South American types, ¢. 9. the Chactinie
and the Bothriuride.

EVIDENCE SUPPLIED BY THE SCORPIONS FOR A LAND-CONNECTION BETWEEN
Souto AMERICA AND AUSTRALIA.

Aithough the following fact has no direct connection with the Scorpion fauna of
Central America, it is sufficiently interesting to call for mention, since it strongly

* In my paper on the Scorpions of the West Indies (Journ. Linn. Soc., Zool. xxiv. pp. 374-404, 1893),
the genera Hadrurus and Brachistosternus were recorded from these islands. The specimens of Hadrurus
and Brachistcsternus were certainly wrongly labelled. The former, in the Berlin Museum, is, as Kraepelin
has shown, an Hadruroides from Ecuador; and the specimen of Brachistosternus, one of the Bothriuride, is
identical with a well-known Peruvian species,

ARACHNIDA. 125

confirms evidence supplied by other groups for a direct land-connection by way of the
Antarctic or the South Pacific between South America and Australia.

‘The family of the Bothriuride is represented in South America by many genera and
species, occurring mainly in Patagonia, Chili, the Argentine, Buenos Ayres, and Brazil.
Beyond the limits of South America the family is unknown, save fur the occurrence
of the single genus and species, Cercophonius squama, which is found all over
Australia, apart from its northern portions, and in Tasmania. Since South America
is obviously the headquarters of the family, it may be inferred that this element of
the fauna of Australia was received directly from South America; but since this
Scorpion, like all Scorpions, is absent from New Zealand, it may be further inferred
that New Zealand formed no part of the land connecting Australia and South
America at the time the ancestor of the Scorpion in question migrated from
South America.

The Scorpions thus supply evidence for a transatlantic connection between tropical
Africa and South America and for a transpacific connection between South America
and Australia: by means of the former America may have received certain elements
of its Scorpion fauna, by means of the latter it may have contributed something to
the fauna of Australia.

Order PEDIPALPI.

The existing Pedipalpi of the suborder Urotricha are referred to a single family,
the Thelyphonide, which extends in the Old World, where it is represented by many
genera, from India to Austro-Malaysia and northwards to China and the Philippine
Islands; and in America from the Southern States of the Union to Brazil. ‘Two
genera only are known from America, Thelyphonellus from the Amazons and Mastigo-
proctus. The latter is represented by a few species in Brazil, a few in the Lesser and
Greater Antilles and two in Central America, one recorded from Guatemala, the other
from Mexico (Cordova, Jalisco, Guerrero, etc.). The Mexican species extends into
Texas and adjoining States of North America.

The nearest ally of Mastigoproctus appears to be the eastern Himalayan genus
Uroproctus. This fact, coupled with the occurrence of the family in China, points to
its former existence throughout the countries bordering the North Pacific, when tropical
or warm temperate conditions prevailed there. If this be so, the tropical American
forms must be regarded as immigrants from the north.

One point connected with the distribution of the group in Central America is its
apparent absence from the States to the south of Guatemala. Nevertheless. it occurs
in the Greater and Lesser Antilles (Haiti, Martinique) and in Brazil. Possibly
therefore it made its way from Central into South America by means of an Antillean

route.

126 ORIGIN, ETC., OF THE FAUNA.

The Amblypygous Pedipalpi have a wider range than the Urotricha. In the Old
World they are represented by two families, the Tarantulide ( =Phrynichide) which
range from Siam, through India, and Arabia into tropical and South Africa, but are
absent from Madagascar, and the Charontide which have their headquarters in
Further India, whence they spread westwards to the Seychelles and eastward to the
Solomon Islands. Like the Urotricha they are absent from Australia *.

The suborder is represented in tropical and subtropical America by the family
Phrynide, which is unknown in the Old World. Of the two subfamilies, the
Heterophrynine are essentially South American, but are unknown in the Antilles.
One species, however, which for want of accurate determination I have provisionally
referred to the Demeraran H. chiracanthus, has been recorded from Central America
without further particulars as to locality.

The Phrynine have a much wider range, extending from North America, throughout
Central America and the West Indies, into South America, Acanthophrynus is known
only from California and Mexico (Durango). Of Phrynus, one species ranges from
Texas to Nicaragua; two occur in Guatemala and are not known elsewhere, and one in
Panama, the latter being found also in Colombia, Venezuela, Trinidad, and Barbados.
Other species occur in the Lesser and Greater Antilles and the northern part of South
America. The third genus, Hemiphrynus, ranges from Mexico (Tuxtla, Oaxaca, Teapa,
etc.), through Guatemala, Costa Rica, and Panama into South America, Although
unknown in the Antilles, there is an outlying species in the Bahamas.

CoNCLUSION.

The Pedipalpi supply no evidence of a mid-Atlantic land-connection between Africa
and America, and none of the existence of a continuous or partial Antarctic land
joining the southern continents. Nor in the case of America do they help much in the
definition of the regions and subregions indicated by the Scorpions. One or two points
may, however, be touched upon.

The occurrences of Acanthophrynus in California and Mexico, of Phrynus whitet in
Texas and Mexico, of Mastigoproctus giganteus } in several of the Southern States of
America and all over Mexico, affiliate the latter country faunistically with the southern
portions of North America. The Sonoran area thus defined differs from South

* One species of Urotricha belonging to the genus Thelyphonus has, however, been recorded from Cape
York, the extreme northern point of Australia.

+ Since the Pedipalpi and Scorpions have existed from the Carboniferous with comparatively slight
structural changes, they cannot be regarded as plastic organisms. In my opinion, therefore, species, genera,
and families have greater significance from the geographical standpoint than groups of those ranks in the
Mammalia.

ARACHNIDA, 127

America in the absence of Heterophrynus and of the Thelyphonid Thelyphonellus ; but
it is linked thereto by the extension of Hemiphrynus from Mexico to Colombia, and of
Phrynus and Mastigoproctus from Texas at least as faras Brazil. The chief distinction
between the Antilles and the Sonoran area is the absence of Acanthophrynus from
those islands, and between the Antilles and South America the absence of Hetero-
phrynus, Thelyphonellus, and Hemiphrynus from the former. But the occurrence of
Phrynus and Mastigoproctus in both the Greater and Lesser Antilles, as well as in

Mexico and South America, points to union between these islands and the countries
in question.

Order SOLIFUGZ.

With one exception the Central American genera of this Order show the same
general geographical features as the Pedipalpi and some of the Scorpions.
Eremobates, the type of a special subfamily, is restricted to the Southern States of
North America and Mexico, its distribution coinciding tolerably closely with that of
the Scorpion Vajovis in the sense that neither extends into the West Indies or into
South America. Ammotrecha, on the other hand, which also occurs in the Southern
States of the Union, is found in the Antilles and in South America, its distribution
being parallel to that of the Pedipalp genera Mastigoproctus and Phrynus and of
the Scorpion Centruroides. The exception above referred to is Hemibdlossia, one
species of which occurs in Guatemala and the others in South Africa. This case of
discontinuous distribution recalls that of the Scorpion Opisthacanthus, whose nearest
allies are tropical and South African species.

Ammotrecha and Hemiblossia belong to a subfamily represented by numerous genera
in Africa, South-western Asia, and the Mediterranean area of the Old World, but
unrepresented in the Oriental Region and Eastern Asia. ‘Their occurrence in Central
America points to an Atlantic connection between the Old and New Worlds. The only
other facts to be borne in mind in this connection are the records of a species of Desia,
typically an African and Mediterranean genus, in Mexico, and of a species of the
genus Lremobates in Afghanistan. Both these records, however, require confirmation,

Order ARANEE.

The Central American fauna of Spiders differs, as a whole, from that of the Scorpions
and Pedipalpi in two particulars, namely, the vast number of species, genera, and
families it is composed of and the large number of these that are common to the Old
and New Worlds. Hence, within the limits of a short essay, it is impossible to deal
with all the families concerned. To illustrate some of the main features of the fauna
and to trace as far as may be its sources, I have selected therefore for detailed treatment

128 ORIGIN, ETC., OF THE FAUNA.

the sharply circumscribed suborder of the Mygalomorphe. The mutual affinities of
the genera and families of this group are sufficiently well understood for the purpose,
and from the geographical standpoint it has the additional advantage of being, so far
as we know, dependent upon continuous land-areas for its dispersal, since the species
appear—for the most part, at all events—to be independent of the method of travelling
commonly known as “ballooning,” which the young of many of the families of
Arachnomorphe have been ascertained to practise shortly after dispersing from the
cocoon. As has been suggested above (p. 119), the very wide, sometimes cosmopolitan
distribution of some genera may perhaps be assigned to this cause, coupled with
exceptional power of adaptation to highly varied conditions.

The Central American representatives of the Mygalomorphe were referred by
Mr. F. Cambridge to the three families, Ctenizide, Dipluride, and Theraphoside.
This classification will be here adhered to in the main as sufficiently exact for the
purpose in hand.

The first subfamily of the Ctenizide, the Actinopodine, is represented by the two
genera, Actinopus and Neocteniza. The latter has only been recorded from Guatemala
and Demerara. The former is abundant all over South America from the Argentine
northwards to Venezuela, and enters Central America at Panama. Perhaps it awaits
discovery in other tropical districts of Central America. It is interesting to record that
_ the only other representative of this subfamily is the Australian genus Missulena
(Eriodon).

The genera of the second subfamily, the Ctenizine, are referred to three sections,
the Pachylomeree, the Ctenizew, and the Cyrtaucheniee. From the first of these
should certainly be dismembered the genus Chorizops as the type of a very special
group, the Cyclocosmee or Halonoproctee. This group contains the three genera:
Chorizops recorded from Mexico (Guanajuato, Vera Cruz), Cyclocosmia from Alabama,
and Halonoproctus from China.

The Pachylomerez comprises three genera: Conothele ranging from Burma to the
Solomon Islands, Hebestatis from California, and Pachylomerus itself. This genus has
a remarkably discontinuous distribution. Several species occur in the Southern States
of North America, in Central America from Mexico southwards to Guatemala and
Costa Rica, and a few in the West Indies (Jamaica, St. Vincent). In the Old World
it js met with in Japan, but, singularly enough, elsewhere only in the western
Mediterranean (Spain, Algeria).

The genus Bothriocyrtum, the sole Central American representative of the Ctenizex,
has one Mexican and one Californian species. Its nearest ally is Cyrtocarenum of the
Mediterranean area. All the other genera of the Ctenizee are Old World forms, two,
in addition to Cyrtocarenum, being Mediterranean, one South African, one Central
Asian, and one Japanese and Chinese,

- ARACHNIDA. | 129

The Cyrtaucheniez, the fourth and last group of the Ctenizine, has three Central
American representatives, Hutychides, Eucteniza, and Enrico, all restricted to Mexico.
One other genus, Myrmekiaphila, from Carolina and Virginia, occurs in the New World.
Of the rest, four in number, one is Burmese, one (Cyrtauchenius) Mediterranean, and
two South African. ‘

The Central American forms of the Dipluride are referred to two subfamilies, the
Diplurine and the Ischnotheline. The Diplurine, corresponding with Simon’s group
Diplurez, have a very singular distribution. In South America there are about nine
genera ranging from Chili northwards, Fufius the single Central American genus
spreading into Costa Rica and Guatemala from Brazil, Ecuador, and Bolivia. Four
genera occur in Australia, two in Madagascar, and one in the Mediterranean area.

The distribution of the Ischnotheline (= Macrothelee, Simon) is equally remarkable.
There are two known Central American genera. vagrus spreads southwards from
Mexico and Guatemala to Bogota, and northwards into the United States (Idaho).
One species, possibly generically distinct from it, has been recorded from South Africa.
The nearest ally of Evagrus appears to be the Transcaspian genus Phyxioschema, which
is itself closely akin to Stenygrocercus from Australia and New Caledonia. Other less
closely related genera occur in the Old World in the Mediterranean area, in tropical
West Africa, Burma, China, and New Zealand. Jschnothele is a very distinct type
from those above mentioned. It occurs in Mexico, Nicaragua, Guatemala, and the
northern countries of South America, and in the Old World, in tropical Africa,
Madagascar, and India.

The Central American genera of Theraphoside * have been revised since
Mr. Cambridge described them. ‘The species he referred to Eurypelma have been
split up into a number of genera and are probably susceptible of further subdivision f.
The species named emilia, pallidum, smithii, and vagans belong to Brachypelma,
which ranges from Mexico to Colombia in South America. ‘Those named ruséicum
ahd seemani are referred to Aphonopelma, which extends from the Southern States of
North America into Mexico and Costa Rica. The species described as Lurypelma
longipes from Guatemala is the only known form of the genus Citharacanthus, which
is allied to Cyrtopholis, a genus mainly confined -to the Greater and Lesser Antilles,
but represented in Central America by C. pernia, referred by Cambridge to Hapalopus,
from Orizaba in Mexico. ‘The genus Dugesiella, containing species from Texas and
Mexico (Guanajuato) has been added to the Central American fauna since

* The Spiders of this family are those commonly referred to in textbooks and Natural Histories as
Mygale, or * Bird-eating ” Spiders.

+ One of the species, spatulatum, was included owing to an error in locality. It belongs to the genus
Grammostolu (== Citharoscelus) and came from Chili. Another Chilian species that was entered and must

‘also be eliminated is Paraphysa manicata.
-- BIOL. CENTR.-AMER., Introd. Vol., June 1915. = S

130 ORIGIN, ETC., OF THE FAUNA.

Mr. Cambridge finished his account of the Spiders. So, too, has the genus Psalmopeus,
which contains the Costa Rican species reduncus and others from Ecuador, Colombia,
and Trinidad. | :

The genera Spherobothria from Costa Rica, Sericopelma from Panama, and
Schizopelma from Mexico and Costa Rica have not, so far, been discovered beyond
the limits of Central America, but, like the others cited above, and the remainder
recorded by Mr. Cambridge, they are related to South American or West Indian
genera of this great family, and with one or two exceptions have no near kinship with
the genera characteristic of tropical Africa, Asia, and Australia, which belong to distinct
groups or subfamilies, Two apparent exceptions to this, however, must be mentioned.
The South American genus Avicularia which extends into Panama belongs to a group
to which also belongs the tropical West African genera Scodra and [Heteroscodra ; and
the genus Hemirrhagus, represented by one species, cervinus, in Mexico, seems to have
its nearest ally in Cratorrhagus of the Mediterranean area.

The conclusion suggested by the above-stated facts is this :—

The Central American Mygalomorphe belong to two categories: (1) Those referable
to groups which are in the main northern in distribution ; (2) those referable to groups
which are in the main southern in distribution.

1. To the northern group belong Pachylomerus, which occurs elsewhere in the
West Indies, North America, Japan, and Spain; Chorizops, whose only known allies
are found in North America and China; Bothriocyrtum, which occurs also in North
America and is related to genera from Japan, China, Central Asia, and the Mediter-
ranean area; Hnrico, Eutychides, Kucteniza, akin to genera from North America,
Burma, and the Mediterranean; Hvagrus, ranging from North America to Bogota,
is allied to genera from the Mediterranean region, West Africa, Transcaspia, China,
and Burma. In the New World none of these groups is found south of the equator.
In the Old World, however, genera akin to Bothriocyrtum, Enrico, Eutychides,
Eucteniza, and Evagrus occur in South Africa, and others allied to Evagrus in
Australia and New Zealand. Perhaps in the future these may be geographically
linked with the northern types, but so far as our knowledge goes they are isolated.

2. To the southern groups belong Neocteniza, from Demerara and Guatemala, and
its ally Actinopus, which ranges from the Argentine into the West Indies and Panama
and is related to the Australian genus Missulena, but to no other genus of the
Old World; Fufius, extending from Brazil to Costa Rica and Guatemala, is related
to many genera confined to South America and to others occurring in Madagascar and
Australia, with one outlying form in the Mediterranean area; Ischnothele, ranging
from the northern countries of South America into Mexico, is also met with in
tropical Africa, Madagascar, and India; all the Central American genera of Thera-
phoside, some of which pass into the Southern States of North America, have their
nearest allies in South America, where they occur at least as far south as the

ARACHNIDA. 131

Argentine and Chili. In the Old World their nearest allies, few in number, are
found in West Africa and the Mediterranean.

Tentative explanations of these facts may be illustrated by one or two cases. If the
groups to which Chorizops and Pachylomerus belong are North American in origin,
they may have extended thence southwards into Central America and northwards into
eastern Asia. Pachylomerus, in addition, may have passed eastwards from America to
Spain and Algeria, or its apparent restriction in the Old World at the present time
to such widely sundered areas as the south-western Mediterranean and eastern Asia
may be attributed to its former extension over Eurasia and its survival only in those
two localities. Against this hypothesis can only be urged the impossibility of assigning
any cause for its extinction in other parts of the Asiatic district.

If the group to which Actinopus and LEucteniza belong was evolved in South
America, these genera may have moved northwards into Central America; and the
restriction of an allied genus to Australia may be assigned to the entry of the group
into that country from South America. |

Since tropical America is the headquarters of [schnothele, judging from the number
of indigenous species, it may be provisionally regarded as the original home of the
genus. Thence it may have passed by a transatlantic bridge to Africa, and so to
Madagascar and India.

These four genera of Mygalomorphe, apart from others which might have been
cited, supply evidence, such as it is, for direct land-continuity, under tropical or
warm temperate conditions, between (1) North America and eastern Asia; (2) North,
Central, or South America and the Mediterranean ; (3) South America and tropical
Africa; (4) South America and Australia.

Corroborative evidence for “Antarctica” is supplied by another family (or sub-
family) of Mygalomorphe, namely the Migide, which is related to the Ctenizide,
but does not occur in Central America. It contains eight genera, distributed as
follows :—Moggridgea and Pecilomigas, South and South-east Africa; yrtale,
Micromesomma, and Thyropeus, Madagascar ; Heteromigas, Tasmania; Migas, New
Zealand; and Calathotarsus, Chili.

It appears to me to be impossible to assign any reason for the extermination of this
group of Spiders in the northern portions of the continents to which they are now
restricted, if the genera in question are the descendants of a family evolved sufficiently
far to the north to have passed from the Old to the New World, or vice versd, along a
North Atlantic or North Pacific route. |

Turning to the Arachnomorphe, one or two of the Central American families may
be selected to illustrate the restricted range of some and the wide range of others.

The Senoculide and Acanthoctenide, each represented by a single genus, are
peculiar to America, ranging from South America northwards to Mexico. ‘The

Psechride are represented by two genera, Psechrus and Fecenia, in the southern and
$2

132 ORIGIN, ETC., OF THE FAUNA.

south-eastern parts of ‘Asia, an area to which they were believed to be restricted until
the discovery of the genus Metafecenia in Central America (Guerrero in Mexico).
The Deinopide (Deinopis), occurring in North, Central, and South America, is also
found in tropical Africa, Madagascar, and South Asia. Of greater interest, because of
their restricted range, are the families Sicariide and Caponiide, both very sharply
defined groups, the latter being also in some respects highly specialized or progressive.
The Sicariide range in America from Chile up the western portions of the continent
as far as Costa Rica, and are only found elsewhere in the world in South Africa.
Similarly with the Caponiide. Of the three genera, Mops occurs in South America
and the Antilles, Caponina in the Antilles and Central America (Guatemala), while
the related genus Caponia is South African. These families corroborate the evidence
derived from other sources for a direct connection between Africa and South America.

The most prominent Spiders in the Central American fauna are those belonging to
the web-spinning species of the family Argyopide. The distribution of some of the
dominant genera of this family is as follows :—Nephila ranges from the warmer States
of North America through:Central America into South America, and is abundant all
over tropical Africa, Asia, and Australia ; Gasteracantha, a hard-shelled spine-bearing
genus, has nearly the same distribution as Nephila. 'The other hard-shelled - spiny
genus, Micrathena, is, however, restricted to America, where its range closely coincides
with that of the two just mentioned. Argyope agrees closely with Nephila, but has
a more northern distribution, at all events, in the Old World, where it reaches the
Mediterranéan area. Finally, the genus Aranea (= Epeira) is practically cosmo-
politan, extending from north temperate, if not subarctic, latitudes southwards all
over the continents.

These genera illustrate, as clearly as others which might have been selected, the
wide distribution of a large number of the genera of Arachnomorphe recorded from
Central America.

CHILOPODA. 133

CHILOPODA, DIPLOPODA, AND PROTOTRACHEATA.
By R. I. Pococg, F.R.S.

The Chilopoda and Diplopoda were in existence in the Carboniferous Period in
Europe and North America. But since these ancient forms were different from their
modern representatives, and since they afford no help to an understanding of the
present distribution of these classes, it is needless to discuss them further, and useless
to repeat what has already been said on this head in the introduction to the essay on
the Arachnida.

Class CHILOPODA (Centipedes).

From the geographical standpoint the Chilopoda or Centipedes are remarkable for
the wide distribution of the orders and of most of the families into which the class is
divisible. It is probable, however, that when the group has been studied in greater
detail, the number of families, especially of the Geophilomorpha, will be considerably
increased and their distribution better understood. The recorded Central American
genera of Geophilomorpha and of Scutigeromorpha call for no special comment in the
present connection, since they exhibit no special features throwing light upon the
origin of the fauna of that country. The Lithobiomorpha and Scolopendromorpha are
more interesting.

The Central American species of the former order belong to the family Lithobiide,
which, although of wide distribution with a few outlying species of the genus Lithobius
in the Southern Hemisphere (India, Australia), is mainly northern, being spread all
over Europe, Central Asia, and North America, and represented by a vast number of
species in these continents. Hence it seems that the Central American representatives
spreading through Mexico to Guatemala must be regarded as a northern element in
the fauna of our area.

The classification of the Scolopendromorpha proposed in my report may be revised
as follows :—

Family Cryproripa*.
Subfamily Cryptopine (Cryptops +, Theatops, Otocryptops, Scolopocryptops).
Subfamily Newportiinz (Newportia, Scolopendrides).
Family ScoLopenpRiIp& (Cupipes t, Rhombocephalus, Scolopendra, Parotostigmus, Rhysida f).

# Brélemann’s examination of the type of Scolopendropsis, showing that the genus is closely related to
Pithopus, proves that the presence of 23 pairs of legs cannot be regarded as a character upon which a group
of family rank can be based. I therefore follow Kraepelin in associating Otocryptops and Scolopocryptops
with Cryptops.

+ Added by Kraepelin to the Central-American fauna since my report was written.

+ In Kraepelin’s Monograph of the Scolopendride (Jahrb. Hamb. Wissen. Arch. xx. 1903, p. 139), this
genus, as expanded by that author, should have been named Trematoptychus, which antedates Rhysida by

one year.

134 ORIGIN, ETC., OF THE FAUNA.

Of the family Cryptopide the genus Cryptops is cosmopolitan, and its discovery in
Central America has no particular significance. Zheatops, Otocryptops, and Scolopo-
cryptops are more restricted in range. Theatops spreads from North America into
Mexico, and has one species in the Mediterranean area. Both Otocryptops and
Scolopocryptops are South, Central, and North American, Antillean, and Chinese,
Otocryptops also extending in the Old World into the south-eastern portion of the
Oriental Region (Philippines, Celebes, Java, New Guinea, etc.) *.

The Newportiine are peculiar to the New World, being plentiful in South and
Central America and the Antilles and represented by one species in North America.
The two genera referred to the subfamily appear to be specialised forms of Crypto-
pide; and, judging from their present distribution, they were probably evolved in
tropical America.

Two of the genera of Scolopendride demand particular attention—namely, Paroto-
stigmus and Scolopendra. ‘The former is mainly tropical American, occurring in
South and Central America and the Antilles, but a few allied forms inhabit tropical
Africa. Of greater interest are the species to which in 1903 (Nat. Hist. Sokotra,
p. 429) I restricted the term Scolopendrat. This genus, as here limited, ranges from
the Southern States of North America, through Central America and the Antilles, into
the northern parts of South America and the Galapagos Islands. Elsewhere it has
only been recorded from the Canary Islands, the Kameruns, Arabia, Syria, Persia,
Sokotra, and the adjoining island of Abd-el-Kuri. Its distribution, therefore, coincides
very closely with that of the Diplocentrine Scorpions, and is suggestive of a direct
land-connection across the tropical Atlantic between the Old and New Worlds.

The Chilopod fauna of Central America shows marked affinities with that of North
America, Asia, and Europe (Lithobius); with Eastern Asia (Otocryptops, Scolopo-
cryptops) ; with tropical Africa (Parotostigmus) ; and with the Mediterranean area
(Scolopendra, Theatops). But there are no data justifying any conclusion as to the
original home of the genera in question, unless superiority in number of species be
regarded as a criterion on that head. If this somewhat unsafe basis for an opinion
be adopted, the inference is that all the genera, except the northern Lithodius, were
evolved in tropical America, and subsequently entered the continents of the Old World
where they are now found.

* One species of Otocryptops, abundant in South and Central America and the Antilles, has also been
recorded from West Africa, There is no proof, however, that the species was not transported, probably in
connection with the slave trade, as 1 have suggested.

t Nearly all the Central American species belong to this genus. The two aberrant forms, punctiventris and

pygmed, should perhaps be regarded as representing distinct genera and need not be considered in the present
connection.

DIPLOPODA. 13

abs |

Class DIPLOPODA (Millipedes).

The dominant Central American forms of the Calobognatha or suctorial millipedes
belong to the genus Platydesmus of the family Platydesmide. This family also has
representatives in North America, Amurland, Malaysia, and the Mediterranean area
in Europe. It has not been recorded as yet from tropical Africa, India, or Australia.
Hence the known facts of its distribution point to its evolution in the Northern
Hemisphere and to its failure to penetrate into the ancient southern continents.
If the Mediterranean species referred to Platydesmus belong, as is probable, to a
distinct genus, Platydesmus is restricted to Central America (Mexico, Guatemala).

The Chordeumoidea are divisible into two families—the Heterochordeumide,
ranging from India to New Zealand, and the Chordeumide, which are abundant in
North America and Europe. Their range eastward from Europe into Asia has not
been ascertained, but there seems no doubt that they are absent from tropical Africa.

The described Central American species, from Mexico and Guatemala, belong to a
genus (Cleidogona) also existing in North America. No species have been discovered
in South America or in the Antilles. This element, therefore, of the Diplopod fauna
of Central America was no doubt derived from North America.

The Stemmiuloidea are an obscure group, known at present only from tropical Asia
(Ceylon), tropical Africa (Liberia), and tropical America (Colombia, Panama, and
Porto Rico). Probably these Diplopods await discovery elsewhere ; but, so far as is at
present known, they may be described as southern. Perhaps the most interesting
point connected with their distribution is the ascription to the same genus (Dopsiuius)
of the Ceylonese, Liberian, and Porto Rican species. ‘This classification accords
with the theory of the former existence of a tropical] transatlantic connection between
the Greater Antilles and West Africa.

The Central American forms of Iuloidea, Paraiulus, from Mexico and Guatemala,

are closely akin to genera and species now living in North America. The nearest ally
of Paraiulus in the Old World is Mongoliulus, recorded from Corea. Here again
there is evidence for a northern derivation of this element of the Central-American
fauna. :
As an explanation of the occurrence of the suborder in North America and Europe,
it may be suggested that it formerly extended across the Northern Atlantic when
temperate conditions prevailed in Greenland, Iceland, and the now-vanished land-areas
which are believed to have joined these countries and Europe and America in one
continuous tract.

Of the Spirostreptoidea, the single Central American genus of Spirostreptide
(Orthoporus) is mainly South American in distribution ; but it is closely allied to, if
not identical with, millipedes of the same family now found in tropical Africa, but not

136 ORIGIN, ETC., OF THE FAUNA.

apparently in tropical Asia. The most plausible explanation of these facts is that
Orthoporus and its near allies were evolved in a continent embracing and connecting
South America and tropical Africa, and that Orthoporus moved northwards into
Central America after the formation of the isthmus of Panama. In Central America
the genus ranges from Panama to Mexico.

Like Orthoporus, the genus Epinannolene also appears to be South American in its
affinities.

Although the precise geographical range of the two dominant Central American
genera of Spiroboloidea, namely Rhinocricus and Spirobolus, has yet to be ascertained,
one or two facts of interest may be noticed.

Spirobolus, ranging from Mexico to Guatemala, occurs also in the Southern States
of North America and in Eastern Asia as far north as Pekin. It is not abundant in
the West Indies, although recorded from Porto Rico. It is apparently absent from
South America. The evidence, therefore, is in favour of a northern, rather than a
southern, origin for this genus. |

Rhinocricus, on the other hand, is abundant in South America and the West Indies,
but not in North America. In Central America it extends from Panama to Mexico.
It is also the dominant genus of the family Spirobolide in the Oriental Region, but
does not appear to extend in Eastern Asia so far north as Spirodolus. Clearly, there-
fore, it is a more southern type than the latter.

There is at present no agreement amongst systematists as to the number of families
into which the Polydesmoidea should be divided, but, setting aside some of the peculiar
forms referred to below, sufficient is known of the inter-relationship of some of the
other genera to make instructive comparison between the Polydesmoid fauna of
Central America and that of other countries.

The most important of these belong to the families Platyrhachide and Chelo-
desmide.

The two subfamilies of Platyrhachide are represented in the tropical south-eastern
countries of Asia, the Platyrhachine being especially abundant in Malaysia. They are
not represented in tropical Africa. In America they inhabit the northern countries of
South America, particularly Colombia and Ecuador, whence the Platyrhachine extend
northwards, at all events, to Costa Rica, and occur in the Antilles; whereas the
Euryurine have a much more northern extension, passing from Costa Rica, through
Guatemala and Mexico, into the Southern States of North America.

The Chelodesmide are very wide-ranging, but from our present point of view the
most interesting forms are the species of the dominant Central American genus
Rhysodesmus of the subfamily Xystodesmine. This genus is represented by a vast
number of species in the northern districts of Central America, but is apparently
absent in South America and in the Antilles. It extends into North America, and
closely allied, if not generically identical, forms occur also in China,

DIPLOPODA AND PROTOTRACHEATA, 137

The Central-American Diplopods, from the faunistic standpoint, belong to three
categories: (1) genera with northern affinities and probably derived from the north;
(2) genera with southern affinities and probably derived from the south ; (8)
restricted or almost restricted to the country and probably autochthonous.

The principal groups of northern origin are the Platydesmide (Platydesmus), the
Chordeumide (Cleidogona), the Luloidea (Paraiulus); Spirobolus of the group Spiro-
boloidea and Rhysodesmus of the Xystodesmine Polydesmoidea.

To the group with southern affinities and probably of southern origin must be
referred the Spirostreptoidea (Orthoporus and Epinannolene), the Stemmiuloidea
(Stemmiulus); Rhinocricus of the group Spiroboloidea and several of the Poly-
desmoidea, like Ligiodesmus, Oncodesmus (Oniscodesmide), Lophodesmus (Pyrgo-
desmide), Platyrhachus, Amplinus, and others (Platyrhachide). Of these, Orthoporus
and probably Hpinannolene have African affinities; the Stemmiuloidea and Pyrgo-
desmidee also have related genera in tropical Africa and Asia. On the other hand, the
Oniscodesmidez and Platyrhachide are at present known only from South-Eastern Asia
and tropical America. ‘The Oniscodesmide may await discovery in Africa, but the
Platyrhachide appear to be certainly absent from that country. ‘Their restriction to
Indo- and Austro-Malaysia and tropical America is singular. Nevertheless, since their
distribution agrees in a measure, though not in exact detail, with that of the existing
Tapiroid mammals, it may be that they are tropical survivors of families once extending
throughout the countries now bordering the North Pacific Ocean.

genera

Of the peculiar, probably autochthonous, groups, the most interesting are the
Polydesmoidea of the family Spheriodesmide, which appear to be a specialised offshoot
of the Chelodesmide. The family ranges in Central America from Mexico to Costa
Rica, and one of the genera, Cyclodesmus, is also represented in Jamaica. Also
apparently restricted to Central America, with the exception of one form recorded from
California, are the genera of the Chelodesmide belonging to the subfamily Rhacho-
desmine.

Class PROTOTRACHEATA.

No fossil forms of this class are known. On morphological grounds it is believed
to be of great antiquity, older even than the Trilobites, but, as Gadow has pertinently
remarked :—* This genus [ Peripatus] need not date further back than the Cretaceous
to allow us to account for the scattered distribution of its species” *.

Until 1894 all the described species of this Class from South America, South Africa,
and Australia, were referred to the single genus Peripatus, despite the ascertained
existence of important morphological differences between those of the Continents in
question. But since that date, when it was first proposed to give new generic names

* Gadow here ignores the work of Bouvier and others, who refer the Prototracheata to several genera.
BIOL. CENTR.-AMER., Introd. Vol., June 1915. T

138 ORIGIN, ETC., OF THE FAUNA.

to the South African and Australasian groups, and to restrict the original title to the
South American groups, great additions have been made to our knowledge of this
Class, mainly by the discovery of new genera, but partly by a more detailed acquaintance
with species that had been previously described. One practical outcome of these lines
of research has been rapid development of the classification from the point of view of
nomenclature, with the result that there appears to be at least eight well-marked
genera distributed as follows :—

I. South and Central America and West Indies.
Peripatus. Tropical America.
Opisthopatus. Chile.

II. Tropical and South Africa.
Peripatus. Congo.
Opisthopatus. South Africa.
Peripatopsis. South Africa.

III. East Indies.
Typhloperipatus. Abor country, N.E. India.
Eoperipatus. Sumatra, Malay Peninsula.

IV. Australasia (including Australia, New Zealand, New Guinea, etc.).
Paraperipatus. New Guinea, Solomon Islands.
Peripatoides. Australia and New Zealand.
Ooperipatus. Australia and New Zealand.

With exception of Typhloperipatus, described in 1913, these genera were admitted
by Bouvier in his Monograph of 1907-8. For my present purpose it is immaterial
that one of the species referred by Bouvier to Ooperipatus has been recently given
generic rank, under the name Symperipatus by Cockerell, and that the species from
the Congo and Chile, assigned by the French author respectively to Peripatus and
Opisthopatus, have been separated from those genera as Mesopertpatus and Meta-
peripatus by Clark*. The interesting point in this connection is that close
relationship exists, on the one hand, between the tropical African and tropical South
American species and, on the other hand, between the Chilian and one of the South
African forms. Clark’s further proposals regarding the generic and subgeneric
divisions, to which he refers the tropical American species assigned by Bouvier to
Peripatus will be referred to later on.

* Tt will be a nice. controversial question for systematists to settle in the future whether the names
‘* Congo-Peripatus” and ‘‘ Chilio-Peripatus,” proposed by Sedgwick for these same species, shall be admitted
as generic terms or not. Clark, perhaps not wisely, disregarded them, probably because Sedgwick, true to his
colours, expressly disclaimed the intention of putting them forward as generic terms. But since they

were available as properly published names, future disputation would have been avoided by accepting
them.

PROTOTRACHEATA. 139

Three classifications of the genera into families and subfamilies have been put
forward by Evans, Bouvier, and Clark. That of Clark, which is a compromise, with
amplifications, between the other two, will serve to show the mutual relationships
of the genera above mentioned admitted at the present time :—

Family Perrpatip”.
Subfamily Peripatine.
Genus Peripatus (Mesoperipatus). Tropical America and Africa.
Subfamily Eoperipatinze.
Genus Eoperipatus. Sumatra, etc.
Genus Typhloperipatus. N.E. India.

Family Periparopsip&.
Subfamily Peripatopsine.
Genus Peripatopsis. 8. Africa.
Genus Paraperipatus. New Guinea, Solomon Islands.

Subfamily Peripatoidine.
Genus Peripatoides. Australia and New Zealand.
Genus Ooperipatus. Australia and New Zealand.
Genus Opisthopatus (Metaperipatus). Chile, S. Africa.

Attention may be drawn to one or two interesting faunistic points revealed by this

arrangement :—

1. Resemblance, in the case of the Peripatide, between the Malaysian forms and
those of tropical West Africa and of tropical South America. ‘This case is
closely paralleled by that of the dwarf Squirrels, Nanosciuride, amongst the
Mammalia.

2. The restriction of the genera of Peripatoidine to Australia, New Zealand,

Chile, and South Africa.
3. The ascription of the Papuasian genus and one of the South African genera to
the same subfamily, Peripatopsine. ‘These genera, however, are not very

closely allied.

THe CENTRAL AMERICAN SPECIES.

Bouvier pointed out in his Monograph that the tropical American species of
Peripatus fall into two groups, which he named Péripates andicoles and Péripates
caraibes. To the former, restricted to the mountains of Central America and the
northern Andes, Cockerell gave the name Oroperipatus, limiting the term Peripatus
to those found in the West Indies and lower-lying countries of South and Central
America. This genus was subsequently divided by Clark into the following subgenera:—

Plicatoperipatus, known from one species (jamaicensis) from Jamaica ; Macroperipatus,
T2

140 ORIGIN, ETC., OF THE FAUNA.

containing species ranging from Rio Janeiro to Central America ; Epiperipatus, also
containing many species with approximately the same distribution as Macroperipatus ;
and Peripatus (s. s.), restricted to species extending from J amaica through the West
Indies to Venezuela.

Oroperipatus, occurring elsewhere in the Andes of Bolivia, Ecuador, and Colombia,
is represented by two species in Central America, namely O. eisent from Tepic in
Mexico and 0. goudoti from Mexico, without further particulars as to locality,
although, as Bouvier suggested, it probably came from the western mountaius of that
country.

Peripatus has many Central American representatives, namely perrieri, from Vera
Cruz in Mexico, geayi, ranging from Cayenne to Panama, both belonging to the
subgenus Macroperipatus; biolleyi and isthmicola from Costa Rica; nicaraquensis
from Nicaragua; edwardsii, extending from Cayenne and Venezuela to Darien and
Panama, and brasiliensis, extending from Santarem on the Amazons to Panama, these
five being referred to the subgenus Lpiperipatus.

It may be noted that Oroperipatus and Macroperipatus are absent from the
Antilles, and that only one representative of Epiperipatus has hitherto been recorded
from those islands, namely from Grenada, the southernmost of the chain *.
On the other hand, Plicatoperipatus and Peripatus (s.s.) are absent from Central
America; while the latter subgenus, exemplified by peculiar species in Jamaica
and Porto Rico and in most of the islands of the Lesser Antilles, exists also in
Venezuela.

CoNCLUSIONS.

The Prototracheata are at the present time restricted to tropical and south
temperate latitudes. They have become adapted to widely different conditions
so far as temperature and climate are concerned. Their extension in the Southern
Hemisphere about as far as the 45th parallel (Tasmania, New Zealand, Isle of
Chiloe), and their existence at high altitudes within the tropics precludes the belief
that their apparent absence from southern temperate areas in the Northern Hemisphere
is due to inability to maintain themselves to the north of the Tropic of Cancer or
thereabouts. The evidence therefore is in favour of the view that the Class was
southern in origin.

The present distribution of the genera Peripatus (sensu lat.) and of Opisthopatus
(sensu lat.) attests a former union between Africa and South America. The
last-mentioned genus also attests, though less strongly, a union between Australasia

* This instance recalls the occurrence of the Armadillo (Dasypus) in Grenada, alone of all the
Antilles.

PROTOTRACHEATA, 141

and South America, since its affinities appear to lie with the Australasian
Peripatoides.

In the case of the tropical American forms, it seems probable that Central America

and the Antilles independently received their prototracheate faunas from South
America.

Recent LITERATURE.
Bouvier, E, L.—Ann. Sci. Nat. Zool. (9) ii. pp. 1-376 (1905) *, and v. pp. 61-318 (1907).
Sepewicx, A.— Quart. Journ. Micr. Sci. lii. pp. 379-406 (1908).

Crarx, A. H.—Smith. Mise. Coll. lx. no. 17, pp. 1-5 (1913); id. Proc. Biol. Soc. Wash. xxvi.
pp. 15-20 (1913).

* In justice to Mr. F. 0. Pickard Cambridge, it should be explained that in Bouvier’s Monograph, figg. 1
and 2, pl.i., depicting examples of Peripatus simoni from Breves on the Amazons, are reproductions of water-
colour sketches taken from life by Mr. Cambridge, who collected the specimens at that locality. ‘The
sketches were sent, with the specimens, to Bouvier from the British Museum, but apparently without any
indication that Mr. Cambridge was the artist and collector.

142 . ORIGIN, ETC, OF THE FAUNA.

Appendix.

The recently published paper by W. D. Matthew, entitled “Climate and Evolution”
(Ann. New York Acad. Sci. xxiv. pp. 171-318, Feb. 1915), came to hand after the
Mamunalian section of this work was in type and after the sections dealing with
the Arthropoda were written. Two of the main propositions of this thesis have an
important bearing upon suggestions that have been put forward to explain some of the
facts in the distribution both of the Mammals and of the Arthropods :—

1. “The principal lines of migration in later geological epochs have been radial
from Holarctic centres of dispersal.” (P. 172.)

2. “The numerous hypothetical land-bridges in temperate, tropical, and southern
regions, connecting continents now separated by deep oceans, which have
been advocated by various authors, are improbable and unnecessary to
explain geographic distribution. On the contrary, the known facts point
distinctly to a general permanency of continental outlines during the later
epochs of geologic time, provided that the allowance be made for the known
or probable gaps in our knowledge.” (P. 173.)

As regards the Mammalia, mentioned in this part of the ‘Biologia,’ whose distribution
is tentatively assigned to vanished land-bridges, Mr. Matthew holds the following
opinions. The resemblance between Thylacinus and the extinct South American
Sparassodont Marsupials, believed by some American paleontologists to indicate close
affinity, is held by Mr. Matthew to be adaptively convergent (p. 265); and Mr. Tate
Regan has quite independently come to this conclusion (British Antarctic ‘Terra
Nova’ Exped. 1910, Zoology, i. no. 1, pp. 41-438, 1914).

Mr. Matthew thinks that the ancestors of the Australian Marsupials entered
Australia by way of Southern Asia, while those of the South American forms came
down from North America. Mr. Matthew ascribes the absence of fossil Marsupials
from Asia to the “imperfection of the geological record,’ and does not discuss either
their comparative paucity in the Austro-Malayan Islands or their absence to the
west of Wallace’s Line—facts, which in my opinion suggest evolution in Australia,
followed by northward migration into Austro-Malaya. Since the Didelphyide have
survived in South America since the invasion of higher placental types, and have even
successfully penetrated into North America, it seems to me that the disappearance of
the Marsupials from Holarctica and their absence at the present time from Indo-
Malaya can only be provisionally assigned to the unsuccessful competition with more

APPENDIX. 143

highly organised groups. However, since the hypothesis of the South Pacific land-
bridge, so far as mammals are concerned, rests largely upon the alleged closeness of
relationship between Thylacinus and, e. g., Prothylacinus, the question need not be
further discussed until the debated point is settled, if capable of settlement, by the
osteological experts concerned.

The difficulty of the South American Monkeys Mr. Matthew also surmounts by
appealing to convergent evolution (p. 216). That is to say, the resemblances between
the Monkeys of the Old and New Worlds have been independently acquired from
Asiatic and South American Lemurs respectively. |

This possibility was long ago considered, and rejected, by Mr. Beddard, but was
adopted by Dr. Scharff. The supposed South American Lemurs—Mr. Matthew admits
that this group is ‘‘ very doubtfully represented in the early ‘Tertiary formations of the
Argentine ”—must have passed, he thinks, into South America from North America
during the Eocene, if I understand him aright. However that may be, I agree with
Mr. Beddard that it is difficult to believe that the relationship between the Cebide
and Simiide is no closer than Mr. Matthew suggests.

In connection with the Hystricomorpha, Mr. Matthew admits that “ we find serious
difficulties ” (p. 229). After discussing the question, he concludes: “I have been
unable to frame any hypothesis which will fit all the facts of the distribution of this
group, except by assuming that the South American Hystricomorpha..... reached
South America from Africa in the Oligocene by over-sea raft-transportation. ‘This
involves so long a voyage that I hesitate to accept it as a reasonable probability, even
though the winds and currents obviously favor transportation in this direction”
(p. 231). Since this suggestion does not seem to be seriously entertained by its
propounder, it does not call for further comment.

The presence of Manatees in the tropical rivers of West Africa and tropical East
America is explained by the hypothesis of the former existence of the genus up the
eastern and western shores of the Atlantic into the Arctic Ocean, to a point where his
map (p.174) shows the extreme north-eastern corner of Greenland may have been
connected with the north-eastern corner of Europe—that is to say, almost to the pole.
This theory will certainly account for the facts; but, until the genus Trichechus
turns up in far northern Tertiary deposits, there is no direct evidence to support it.

Although Mr. Matthew deals mainly with the Vertebrates, he remarks in connection
with the supposed transatlantic bridge between Africa and South America: “The
supposed evidence in its favour from lower vertebrates and invertebrates is due, so far
as I have been able to examine it, to a lack of appreciation of the principles of
dispersal of races and of parallelism and of the imperfection of the geological record ”
(p. 231). Presumably he holds the same opinion regarding the supposed bridge
between South America and Australia. Nevertheless, I cannot bring myself to believe

144 ORIGIN, ETC., OF THE FAUNA.

that the close likeness between the Australian Scorpion Cercophonius and several
South American genera nor the resemblances between the genera of trap-door Spiders
referred to Migide (p. 131) are due to parallel evolution. Nor is it at all intelligible
to me why the Scorpions and Spiders in question have disappeared entirely from all
parts of the world except the southern continents, if at one time they were Holarctic
in distribution. Nor can I think that raft-transportation will account for their present
discontinuous range. Whether the land-bridge theory is the true explanation is,
of course, quite another matter; but it appears to me to be the least improbable of
the four *.

* Elevations of ocean-floors to connect continents are usually regarded as vast changes. From the human
standpoint the epithet is justifiable; but, if the diameter of the earth be taken as a standard, such changes
raay be described as infinitesimal.

[ 145 ]

FLORA.
By W. Bortine Hemstey, LL.D., F.R.S.

_ Since the publication of the Appendix and Introduction to the “Botany” of the
‘Biologia Centrali-America,’ in 1888, botanical explorers have been very active, not
only within our limits and in the contiguous countries, but also more especially in
Africa and Eastern Asia, from Burma and China, southward and eastward, through the
Malayan Archipelago to the Philippines, New Guinea, and Fiji.

The addition of new species to the Central American Flora is enormous, especially
from Mexico and Guatemala*. Of new genera, established on newly discovered
types, there are relatively few; but very many new genera have been founded by
the segregation of old and familiar genera.

Notable among the discoveries in tropical and subtropical districts are additional
genera and species of southern types or families, belonging to the Vochysiacee,
Trigoniacee, Lecythidacez, Lauracee, Kuphorbiacee, Artocarpacee, etc., etc.

The identification of Schlechtendal’s Mexican genus Juliania, the discovery of
several new species of this genus, and the founding of the family Julianiacew, including
the exceedingly rare and imperfectly known Peruvian monotypical Orthopterygium,
constitute a most interesting botanical contribution.

But, taken on the whole, the internal discoveries throw no further light on phyto-
geography than we possessed in 1888. So little was known of the natural history of
Lower California that it was not included in the ‘ Biologia,’ though politically and
geographically belonging to Mexico. It has since been sufficiently explored, as well
as the outlying islands of Guadalupe, the Revillagigedo Group, and others, to reveal
the characteristics of the vegetation and flora, which are essentially Sonoran, with an
intermixture in the north of Upper Californian species. The endemic element, alike
in the Peninsula and the Islands, is of no greater proportion than that of the adjacent
mainland. American botanists have devoted much time to the investigation of the
vegetation of North Mexico and of the States and Territories north-west of Mexico
with extremely interesting results, emphasising the distinct origin of the Pacific and
Atlantic floras. |

* We have no statistics, except those kindly furnished for Guatemala by Captain John Donnell Smith,
who, in his own publications, has added eighty-four genera and 1224 species not included in the ‘ Biologia.’
Eleven of the genera and 488 of the species are new. It may, however, be safely assumed that about
2000 genera of flowering plants are represented in Central America by at least 15,000 species.—W. B. H.

BIOL. CENTR.-AMER., Introd. Vol., June 1916. | U

146 ORIGIN, ETC., OF THE FLORA.

Outside of America, the exploration of Central Africa has yielded the most startling
results. Contrary to expectation, the relationships between the African and American
floras are almost as strong as those between the African and Asiatic floras, including
representatives of such otherwise exclusively, or almost exclusively, American families
as the Canellacee, Caricaceez, Humiriacee, Hydnoracee, Loasacee, Mayacacee,
Napoleonacez, Rapateacee, Strelitziacee, Velloziaceee, and Vochysiacee. There are
equally striking generic and specific connections. Altogether there are records of the
same, or specially representative, genera and. species belonging to upwards of sixty
families or distinct groups. These facts, coupled with the zoo-geographical data
and the paleontology of the region, have given rise to the theory of a Jand-connection
between Africa and South America in Eocene times, a theory accepted by both
botanists and zoologists of high repute. Von Ihering’s projected map of the assumed
‘“Verteilung von Land und Meer zur Eocién-Zeit ” offers a plausible solution of some
of the problems of the present longitudinal distribution of organisms, alike in southern
latitudes and in remote islands. As long ago as 1885 (Introduction to the Botany of
the ‘Challenger’ Expedition), I rejected the theory of the vegetation of remote volcanic
islands and groups of islands, such as the Galapagos and Hawaii, being necessarily
of derived origin rather than remnants of a former wider flora. This view does not
exclude dispersal by various agencies in distant parts of the world and natural
migrations across the widest continents, as exemplified by some certainly introduced
plants. .

The explorations of the last twenty years in Western and Central China furnish
further evidence of the existence of close relationships between the floras of eastern
temperate Asia and eastern North America. Types illustrating this feature are.
usually representative species of near affinity, as, for example, of the genera
Liriodendron, LIiquidambar, and Sassafras. This American-Asiatic element extends
southward to Mexico, and is more prominent in the mountains of eastern South
Mexico than it isin the north. The following genera of forest-trees represented in
the Atlantic States, absent from the Pacific States, reach Mexico, namely :—Magnolia,
Asimina, Tilia, Robinia, Liquidambar, Ilex, Diospyros, Bumelia, Ulinus, Celtis, Morus,
Ostrya, Carpinus, and Carya. In a general sense, the western coast-forests are
coniferous and the eastern deciduous. Certain genera are apparently now confined,
or nearly so, to America and the Mediterranean region (including the Azores and
Canaries)—such are Lelianthemum, Lupinus, Heberdenia, Platanus, and Corema.

Apart from the subantarctic flora, there are genera that extend from Chili to Mexico
and from Australasia to Borneo. The magnoliaceous genus Drimys belongs to this
category. Judging from analogies, this is an example of a northern extension of
a southern type. The Proteaces, now almost exclusively confined to the Southern
Hemisphere, furnish a similar instance in the closely allied genera Helicia and Roupala.
The former ranges from Australasia to China, Japan, and India, the latter from
Brazil to Mexico, and neither has further extensions. Of course, the absolutists

BOTANY. 147

would regard these as tardy stragglers in a southern migration. The phanerogamic
flora of Central America (as calculated in 1888) comprised an intermixture of northern
and southern types of varying proportions in different areas and an autochthonous
element. The last constituted only eleven per cent. of the genera against seventy
per cent. of the species. The percentages of genera restricted to America and of
wider ranges were 53°7 and 46:3 respectively. For species the figures stand at
89°9 per cent. restricted to America and 10°1 per cent. extending beyond America.

The statistics of a specimen of the upper mountain-flora comprise 260 genera, of
which eighty-two (or 31°6 per cent.) were restricted to America, thirteen (or 5 per
cent.) endemic within our limits, and the rest, 165 (or 63:4 per cent.), had extra-
American extensions.

_ Nine of. the exclusively American genera had northern extensions ; twenty-five had
southern extensions; twenty-two were common to the Andes only; and twenty-six
were also represented in both North and South America.

The total number of species enumerated is 604, whereof 504 (or 83-4 per cent.) were
reckoned as endemic; 83 others (or 13:8 per cent.) not extending beyond America,
with a residue of only 2°8 per cent. of wider range. The extensions to other parts of
America are given as: N. America, 17; 8. America, 19; Andes only, 39; N. and
S. America, 8. Of course, most of these figures are rough approximations, but they
are sufficient to show that the representatives of northern and southern types are
nearly balanced, and that the Andine element in the mountain-flora of Central America
preponderates over the temperate northern element *.

‘Taking the whole phanerogamic flora of Central America, as it was known in 1888,
638 of the genera had north-western extensions, 562 north-eastern connections, whilst
1285 had southern extensions. Species yielded similar proportions. But statistics
alone explain very little. | |

The present complexities of plant-distribution point to greater and oftener repeated
alterations in the distribution of land and water than is generally admitted, and no
one theory is sufficient, in my estimation, to account for the origin and progressive
dispersal of organisms. Indeed, it is doubtful whether sufficient evidence still exists
to carry us to a convincing conclusion. ‘There is always the great question whether
organic as well as inorganic matter has not developed on the same or similar lines in
different regions or centres. Comparing the following particulars of the distribution
of families, it is difficult to realize that one part of the world has produced a land-
vegetation wholly different in composition from:that of any other part. Very diverse
interminglings exist, but they give little or no clue to the beginnings. |
- Accepting, for purposes of comparison, the number of families of flowering plants
at 290, as defined in the seventh edition of Engler’s ‘Syllabus,’ very nearly three-
fourths are represented in both the Eastern and Western Hemispheres. In the more

* Of, Biol. Centr-Am., Botany, iv. p. 299 (1887).
v2

148 ORIGIN, ETC., OF THE FLORA.

restricted sense of Bentham and Hooker, four-fifths are common to both Hemispheres.
The families confined to America are twenty-five in number, namely, Batidacee,
Bromeliacez, Calyceraceee, Cannacez (Scitaminee *), Caryocaraceze (Ternstreemiacee),
Columelliacee, Cyclanthacee, Cyrillacee, Fouquieracee (Tamariscine), Gomortegacee,
Lacistemacee, Leitneriacee, Lennoaceee, Limnanthacee (Geraniacee), Malesherbiacee,
Marcgraviacee (Ternstroemiaceze), Martyniacee (Pedalinee), Myzodendracee, Qui-
inacee, Sarraceniacee, Thurniaceee, Tovariacee (Capparidaceze), and -'Tropzolacez
(Geraniacee).

The foregoing families are of various categories. Eleven of these are limited
to a single genus each, and the Bromeliacee is the only one exhibiting a considerable
development of genera and species, now numbering forty-five and about a thousand
respectively, generally dispersed in tropical and south temperate America. Twelve
out of these twenty-five families are not known to be represented within the limits
of the ‘ Biologia.’

Fourteen families are peculiar to the African region (including Madagascar and the
Mascarene Islands), so far as known, eight of which are monogeneric. The only one
of considerable development is the herbaceous Selaginacez, associated with the
Scrophulariaceee by Engler. The Chlenacee, comprising six genera and about
twenty-five species, are peculiar to Madagascar. ‘They belong to the Malvales, and
are shrubs and small trees with showy flowers.

A group of small and interesting families inhabiting eastern Asia, consisting of the
Cercidiphyllacee, Kucommiaceer, Pentaphylacacee, Stachyuracee, and Trochodendracee,.
of remarkable affinities, is not represented in North America, but Engler places
Pentaphylax near the American Cyrillacez.

In spite of its highly differentiated vegetation, the Australian flora counts few
peculiar families; indeed, the only ones absolutely limited to Australia are the
Cephalotacew, restricted to the singular Cephalotus follicularis, the Eupomatiacee
and the Tremandracee, allied to the Pittosporacew, which, except the widely spread
genus Pittosporum, are all Australian. But there are some characteristic Australasian
(including New Zealand, New Caledonia, etc.) families with a few solitary outliers—
such are the Centrolepidacee, represented by one species in China and one or two
species in the extreme south of America, and the Epacridace, with a few outliers in
Malesia and Polynesia, and the monotypic Lebetanthus in Fuegia. The Eucryphiacee
(Eucryphia), limited to about four species in Australasia and Chili, and the somewhat
numerous and generally dispersed Australian Stylidiacee are represented outside of
Australasia by solitary outliers only in India, Malaya, and Fuegia.

In addition to the above-named families, there are some twenty-five others peculiar
to the Old World (with some extensions in Polynesia), but inhabiting two or more of

* The family-name under which the groups in question appear in the “ Botany ” of the ‘ Biologia.”—W.B.H, -

BOTANY, 149

the great divisions of the Eastern Hemisphere. No fewer than fourteen of these are
monogeneric and inconspicuous in the vegetation, except Casuarina and Nepenthes.
Casuarina and Pandanacee are prominent and widely spread in maritime districts, yet
they have not reached America independently of human agency. ‘The Flagellariacez
and Philydracew also largely affect coastal regions, ‘The arboreous Dipterocarpacew,
numbering some 300 species, is the only large family in this category, and it
forms a conspicuous feature in the forests of India and Malaya—rare in Africa, and
apparently absent from Australia. .

Including the families respectively nearly or quite peculiar to Africa, to Eastern
Asia, and to Australasia, there are fifty-three families in the Old World which are not
represented in America, as against twenty-five restricted to the New World.

From the foregoing particulars of distribution it is evident that harmonies do not
exist on a large scale in the same way that obtains, for example, in the different
islands of the Galapagos Archipelago. ‘The peculiarities of plant-distribution are
infinite and reducible to no laws: take, for illustration, the composition and present
distribution of the families of root-parasites in relation to their affinities—the widely
spread Balanophoraceze and Orobanchacex and the local Lennoacee of North Mexico,
or such essentially insectivorous families as the Nepenthacee of the Old World
and the Sarraceniacez of America and the widely spread Lentibulariacee. One
might go on multiplying the exposition of the anomalies and curiosities or phenomena
of distribution. But just one more example: the Lardizabalacee comprise about
half-a-dozen small genera, ‘several of them distinct monotypes ; two of the genera are
endemic in Chili, and the rest of the order is restricted to North India, China, and
Japan.

Mr. R. I. Pocock, in his account of the Mammalia, seems to have arrived at the
same inconclusive results as myself in respect of the plants; but he evidently favours
a former southern land-connection as a solution of the main problem, and, barring
independent developments of life, it is the only theory that commends itself to my
mind. No other explanation seems adequate *.

* It was originally planned to discuss more fully in this plaee the composition and origin of the Central
American Flora, and at least six months were devoted to the collection of materials for this purpose; but a
combination of adverse circumstances has hitherto hindered the completion of the work and rendered it
impossible within the immediate future. It is a great disappointment to me that I could not furnish
Dr. Godman with something in my line more worthy of his monumental publication.

I have great pleasure in recording the faet that I have received much assistance from American Botanists,
who have most liberally presented their published works. My thanks are more especially due to Prof. T. 8.
Brandegee, Dr. N. L. Britton, Mr. H. Pittier, Dr. B. L. Robinson, Dr. J. N. Rose, Captain J. Donnell Smith,

and Dr. W. Trelease.—W. B. H.

PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET,

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