FOOD AND STORAGE.

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Dipodomys s. spectabilis does not hibernate, so must prepare for unfavorable seasons by extensive storage of food materials. There are two seasons of the year, in southeastern Arizona at least, when storage of food takes place, namely, in spring, during April or May, and in fall, from September to November, the latter being the more important. For the periods between, the animal must rely largely on stored materials. Not infrequently a season of severe drought precludes the possibility of any storage. The summer and fall of 1918 was such a season on the Range Reserve (Pl. II, Fig. 2). If food stores are inadequate at such a time the kangaroo rats must perish in considerable numbers. Fisher found many deserted mounds in the vicinity of Dos Cabezos, Ariz., in June, 1894, which may be accounted for in this way. In 1921 Vorhies found all mounds within 4 or 5 miles of Albuquerque, N. Mex., deserted by spectabilis, resulting probably from overgrazing by sheep and goats during a succession of dry years. In the arid Southwest natural selection probably favors the animals with the largest food stores, and it is not surprising that the storing habit has been developed to a remarkable degree.

Some stored material is likely to be found at any time of year in any mound examined, the largest quantity usually in fall and winter, the smallest in July or August (Table 1, dens 1, 2, 14, and 24). Amounts found by different observers vary from a few ounces to several quarts or pecks, and stored materials taken from 22 mounds on the Range Reserve vary in weight from 5 to 4,127 grams (more than 9 pounds). This is exceeded by one lot from New Mexico, which totaled 5,750 grams (12.67 pounds). It is fairly evident that in seasons of scanty forage for stock the appropriation of such quantities of grass seeds and crowns and other grazing materials by numerous kangaroo rats may appreciably reduce the carrying capacity of the range. Studies of cheek-pouch contents and food stores taken from dens show that the natural food of spectabilis consists principally of various seeds and fruits, particularly the seeds of certain grasses. The study of burrow contents has been especially illuminating and valuable.

All of the stored material from 22 dens on the Range Reserve and from 2 near Albuquerque, N. Mex., has been saved and analyzed as to species as carefully as the conditions of storage would permit. Within the mound the food stored is usually more or less segregated by plant species, though the stores of material of any one kind may be found in several places through the mound, and often the material is mixed. In the latter case the quantities of the various species can only be estimated, but in the former the species may be kept separate by the use of several bags for collecting the seeds, and a fairly accurate laboratory weighing can be made later. Very frequently, the explanation of this separation of species lies in the different seasons of ripening, but sometimes where two species are ripe at the same time near the mound, one is worked upon for a time to the exclusion of the other. The one kind is often packed in tightly against the other, but with a very abrupt change in the character of the material.A number of the more interesting and representative results of the weighing and analyses of burrow contents are presented herewith in tabular form. The data for each den, or lot, shows in grams the quantity of stored material removed and the best estimate it was possible to make of the percentages or weights of the various species. When the weight was less than 5 grams, the mere trace of the species frequently is indicated in the following tables by the abbreviation "Tr."


Table 1.Analyses of plants stored by Dipodomys spectabilis spectabilis Merriam, obtained from examination of representative dens (all except Den 24 from U.S. Range Reserve, near the Santa Rita Mountains, Ariz.).

Den 1.

February 7, 1918. Burrow typical, located on bank of wash in partially denuded grass-land, Bouteloua rothrockii and weed type; soil sandy; burrow photographed in section (Pl. VII, Fig. 1).

Species stored. Grams.
Bouteloua rothrockii 2,205
Bouteloua aristidoides (B. eriopoda and B. rothrockii, Tr.) 1,445
Plantago ignota 442
Eriogonum polycladon 35

Total 4,127

Four species of plants represented in burrow contents (Pl. VII, Fig. 2). Maximum quantity for single burrow in series of 22 from Range Reserve.

Den 2.

March 9, 1918. Surroundings overgrazed and partially restored by complete protection. Red soil, with much coarse rough gravel and stone.

Species stored. Grams.
Bouteloua rothrockii (nearly pure) 1,460
Bouteloua rothrockii (mixed with Aristida spp.) 945
Boerhaavia wrightii 660
Bouteloua rothrockii } 525
Bouteloua aristidoides
Aristida divaricata
Aristida bromoides
Kallstroemia laetevirens Tr.
Heterotheca subaxillaris Tr.
Plantago ignota 15
Fleshy fungi 10

Total 3,615

Eight species of plants represented by seeds. One species of fleshy fungus in addition.

Den 4.

September 20, 1918. In Calliandra type. Stony or gravelly soil, red, nearly denuded of grass.

Species stored. Grams.
Prosopis velutina 190
Mollugo verticillata (pure) 90
Anisolotus trispermus (mixed, but mostly of this genus) 50
Solanum elaeagnifolium (12 fruits) 2
Per cent.
Mollugo verticillata (inseparable) 50 } 400
Bouteloua rothrockii 1
Bouteloua aristidoides 10
Lepidium lasiocarpum Tr.
Polygala puberula Tr.
Ayenia microphylla 2
Portulaca suffrutescens 1
Aplopappus gracilis Tr.
Alternanthera repens 1
Tridens pulchella 1
Plantago ignota 33
Panicum hallii Tr.
Fleshy fungi (puffballs) 2

Total 734

Fifteen species represented in addition to the fleshy fungi. No perceptible grass growth from the summer rains here, therefore dependent on a wide variety of scattering plants.

Den 6.

October 17, 1918. Mixed type, partially denuded, no growth from summer rains. Sandy soil.

Species stored. Grams.
Bouteloua rothrockii (crowns) (heads 1 to 2 per cent) 1,435
Bouteloua rothrockii (heads and crowns, about 50 per cent of each) 325
Bouteloua rothrockii (with small percentage of crowns) 315
Boerhaavia wrightii (with a few grass crowns) 150
Prosopis velutina 90
Solanum elaeagnifolium (3 fruits) Tr.

Total 2,315

Four species represented. Count of 100 grams of stored Bouteloua crowns gives 1,700, or 17 crowns per gram. At this rate there were at least 27,000 crowns stored in this burrow. If a density of 250 plants to the square yard be assumed (a high estimate) these crowns represent the total B. rothrockii on 104 square yards of range surface. Further examination of the vicinity of this den showed that the surrounding area was not completely cleared, but was devoid of B. rothrockii, while still having B. eriopoda with crowns undisturbed.

Den 11.

April 9, 1919. In partially denuded land where good spring growth of Eschscholtzia was in bloom at time of excavation. Stomach of spectabilis killed in this burrow contained a mass of fresh but finely comminuted green material, probably poppy leaves, strongly colored with yellow from blossoms. No summer growth here in 1918.

Species stored. Grams.
Bouteloua rothrockii (crowns) (miscellaneous chaff, etc.) 107
Eschscholtzia mexicana (buds and flowers) } 10
Anisolotus trispermus (leaves and pods)
Gaertneria tenuifolia (leaves)
Lupinus sparsiflorus (flowers)
Solanum elaeagnifolium (2 fruits) Tr.

Total 117

Six species represented, some only by leaves or flowers and not by seeds. Such storage is never in large quantity. The fresh storage material was weighed after becoming air dry. This illustrates a late spring condition, storage running low.

Den 14.

August 8, 1919. Excellent summer growth all over range. This burrow in mixed growth, grasses and weeds.

Species stored. Grams.
Miscellaneous portions of green plants of mixed species, no seeds 5

Representing minimum for any one of the 22 burrows studied. Active storage does not begin until September.

Den 16.

October 17, 1919. In good grass, but mound overrun by a large Apodanthera vine.

Species stored. Per cent. Grams.
Aristida divaricata 90 to 95 } 58
Chamaecrista leptadenia 10 to 5
Bouteloua rothrockii Tr.
Prosopis velutina 200
Apodanthera undulata 55

Total 313

Five species represented. Two species, Apodanthera, and Chamaecrista leptadenia, new to storage records. Several whole fruits of Apodanthera, about 2 inches in diameter, stored in addition to seeds alone; seeds of this form not previously noted in burrows, but very abundant in this one, indicating importance of the factor of accessibility in storage.

Den 19.

October 31, November 1, 1919. In good grass. Entire burrow system mapped (Fig. 2, p. 29).

Species stored. Per cent. Grams.
Aristida spp. (probably mostly divaricata) 98 } 1,813
Eriogonum sp Tr.
Bouteloua rothrockii 1
Bouteloua aristidoides 1
Panicum sp Tr.
Prosopis velutina 1,213

Total 3,026

Five species represented, in addition to those of Aristida. Largest storage of Prosopis found. Mound was near a good-sized mesquite tree. No storage in subsidiary burrows.

Den 21.

January 31, 1920. Male trapped here night of January 29, and suckling female trapped at same place and same opening of mound, night of January 30. Burrow excavated to secure young, which were found in special nest chamber.

Species stored. Grams.
Aristida spp. (intimate mixture of undetermined species) 1,115
Eschscholtzia mexicana (from spring of 1919) 48
Opuntia (prickly pear, seeds only, no fruits) 10

Total 1,173

Three species represented. Prickly pear hitherto found as fruits only.

Den 22.

January 1, 1921. Rather good grass growth here in summer of 1920. Burrow typical, sandy soil. Two skulls of former residents unearthed.

Species stored. Grams.
Aplopappus gracilis (some B. rothrockii) 1,030
Astragalus nuttallianus 630
Bouteloua rothrockii (some A. gracilis) 530
Sida diffusa 30
Solanum elaeagnifolium (282 fruits) 53
Loeflingia pusilla Tr.
Bouteloua aristidoides Tr.
Plantago ignota Tr.
Lupinus sparsiflorus Tr.
Old storage (mostly Bouteloua aristidoides with traces of B. rothrockii and Aristida divaricata) 60

Total 2,333

Eleven species represented. First instance of quantity storage of Aplopappus gracilis. First occurrence of Loeflingia pusilla and Astragalus nuttallianus.

Den 24.

November 8, 1921. On mesa northeast of Albuquerque, N. Mex., near base of Sandia Mountains. Fair grass growth here during preceding summer.

Species stored. Grams.
Sporobolus cryptandrus strictus 5,455
Salsola pestifer 295

Total 5,750

Two species represented. The heads of Sporobolus cryptandrus strictus are retained to a great extent within the leaf sheaths. This necessitates the cutting of the stems into suitable lengths for carrying, and the stored material appears to be merely cut sections of the stems. Close examination, however, discloses the heads within, and shows that as in other instances seed storage is the end sought. These pieces are packed beautifully parallel like so many matches, and vary from a minimum length of 20 to a maximum of 37 millimeters, averaging about 30. Count of 2 grams of the above Sporobolus material shows that there are 125 separate cut sections per gram, or a total of approximately 680,000 pieces in this one lot of storage, indicating a remarkable activity on the part of the individual rat (Pl. VIII, Fig. 1).

Plate VII. Fig. 1.—Den Excavated on Range Reserve. Plate VII. Fig. 1.—Den Excavated on Range Reserve.
Vertical section through Den No. 1, of Table 1 (p. 20), showing the complex system of burrows, some of them plugged with closely packed storage (outlined in white), the depth of the den, and the widened chambers centrally located.
Plate VII. Fig. 2.—Content of Excavated Den. Plate VII. Fig. 2.—Content of Excavated Den.
Storage content of Den No. 1 (Fig. 1, above), showing the separate species of plants listed in Table 1. The rod is 1 meter long. The large pile on the left is composed of seed-laden heads of crowfoot grama (Bouteloua rothrockii), the large pile on the right consists of heads of six-weeks grama (Bouteloua aristidoides), the pile of heads in the center is desert plantain (Plantago ignota), and the smallest heap is composed of buckwheat-bush seeds (Eriogonum polycladon).

The number of lots of storage (24) studied in detail, extending as it does over a period of three years with seasons of varying growth conditions, is not sufficient to permit the construction of a curve showing increase and decrease in quantity of stored material with growing seasons and intervals between; but the results indicate a very decided increase during the autumn storing season, and continuing large well into the winter, since some outside material can still be obtained until midwinter. From about February to April a decrease may be noted, followed, if the spring growth of annuals be good, by a slight increase; and we can very nearly predict the general character of the increases and decreases by the precipitation and consequent growth conditions.

Table 2.Quantity of storage per den correlated with time of year and growth conditions of preceding season (chiefly from United States Range Reserve near the Santa Rita Mountains, Ariz.).

Den No. Date. Quantity. Preceding season.
1918. Grams.
1 Feb. 7 4,127 Good.
2 Mar. 9 3,615 Do.
3 July 25 401 Poor.
4 Sept. 20 734 Do.
5 Sept. 21 2,520 Do.
6 Oct. 17 2,315 Do.
7 Dec. 20 1,247 Do.
1919.
8 Feb. 7 1,600 Do.
9 Mar. 13 370 Do.
10 Apr. 7 180 Do.[3]
11 Apr. 9 117 Good.[3]
12 May 7 298 Do.[3]
13 May 11 1,590 Do.
14 Aug. 8 5 Good.
15 Sept. 4 151 Do.
16 Oct. 17 313 Do.
17 Oct. 18 583 Do.
18 Oct. 25 3,410 Do.
19 Nov. 1 3,026 Do.
20 Dec. 13 2,816 Do.
1920.
21 Jan. 31 1,173 Do.
1921.
22 Jan. 1 2,333 Fair.
23[4] Nov. 7 1,685 Good.
24[4] Nov. 8 5,750 Do.

In presenting Table 2, showing quantity of storage per burrow correlated with the time of year and the character of the preceding growing season, the fact may be emphasized that the growing seasons in southern Arizona are two in number—early spring and midsummer. The spring season is the less important, the plants consisting chiefly of a variety of small annuals, while the important range grasses make their chief growth and head out almost exclusively in the July-August rainy season. It may be noted also that the actual increases in storage appear somewhat after the growth period proper, since storing does not get well under way until the seed crop is mature. The banner-tailed kangaroo rat shows a marked adaptability to different foods available in the neighborhood of its burrows. It must, perforce, adapt itself and its storage program to the food that it can get, and this varies enormously with the climatic conditions of successive seasons. The large numbers present in suitable localities clearly indicate that the animal is successful in meeting the changing and sometimes extremely adverse conditions of its environment.

Plate VIII. Fig. 1.—Content of Den Excavated in New Mexico. Plate VIII. Fig. 1.—Content of Den Excavated in New Mexico.
Storage content of Den No. 24, of Table 1, from Sandia Mountains, N. Mex. This is the largest lot of storage taken in the course of the investigations. The larger pile consists wholly of a valuable grass, Sporobolus cryptandrus strictus: the smaller of Russian thistle (Salsola pestifer.)
Plate VIII. Fig. 2.—Growth Following Elimination of Kangaroo Rats. Plate VIII. Fig. 2.—Growth Following Elimination of Kangaroo Rats.
The same mound as shown in Plate III, Figure 1, after three years of protection, the rodents having been killed out. Nearly as good grass recovery following poisoning operations occurred in the single excellent season of 1921.

At times, more especially in the seasons of active growth, some of the green and succulent portions of plants are eaten. This was very noticeable in the spring of 1919, when a most luxuriant growth of Mexican poppy (Eschscholtzia mexicana) occurred. Stomachs at this time were filled with the yellow and green mixture undoubtedly produced by the grinding up of the buds and flowers of this plant. Small caches of about a tablespoonful of these buds were also found in the burrows at this time. Occasionally in spring one may find a few green leaves of various plants, Gaertneria very commonly, tucked away in small pockets along the underground tunnels, indicating that such materials are used to some extent. As has been shown in detail, however (Table 1), the chief storage, and undoubtedly the chief food, consists of air-dry seeds.

The character of the storage, the absence of rain for months at a time in some years, and the consequent failure of green succulents show that without doubt spectabilis possesses remarkable power, as to its water requirements, of existing largely if not wholly upon the water derived from air-dry starchy foods, i.e., metabolic water serves it in lieu of drink (Nelson, 1918, 400), this being formed in considerable quantities by oxidation of carbohydrates and fats (Babcock, 1912, 159, 170). During the long dry periods characteristic of southern Arizona, no evidence that the animal seeks a supply of succulent food, as cactus, is found; and if it may go for two, three, or six months without water or succulent food, it is reasonable to suppose that it may do so indefinitely. In the laboratory spectabilis ordinarily does not drink, but rather shows a dislike for getting its nose wet. During the periods of drought the attacks upon the cactuses by other rodents of the same region, as Lepus, Sylvilagus, Neotoma, and Ammospermophilus, become increasingly evident. The list of plant species thus far found represented in the storage materials of spectabilis on the Range Reserve is shown in Table 3.

Table 3.List of all plant species found in 22 dens of Dipodomys spectabilis on the United States Range Reserve, near the Santa Rita Mountains, Ariz., with approximate total weights.

Grasses.
Grams.
Aristida bromoides (six-weeks needlegrass) 536
Aristida divaricata (Humboldt needlegrass) 9,412
Aristida scabra (rough needlegrass) 344
Bouteloua aristidoides (six-weeks grama) 3,093
Bouteloua radicosa (grama) 1,269
Bouteloua eriopoda (black grama) Tr.
Bouteloua rothrockii (seeds, 8,495; crowns, 3,517 grams) (crowfoot grama) 12,012
Festuca octoflora (fescue grass) 70
Panicum arizonicum (Arizona panic-grass) 11
Panicum hallii (Hall panic-grass) Tr.
Pappaphorum wrightii Tr.
Tridens pulchella Tr.
Valota saccharata Tr.
Other Plants.
Alternanthera repens Tr.
Anisolotus trispermus (bird's-foot trefoil) 186
Aplopappus gracilis 1,030
Apodanthera undulata (melon loco) 55
Astragalus nuttallianus (milk vetch) 630
Ayenia microphylla Tr.
Boerhaavia wrightii 885
Chamaecrista leptadenia (partridge pea) 5
Echinocactus wislizeni (visnaga) 5
Eriogonum polycladon 35
Eschscholtzia mexicana (Mexican poppy) 250
Gaertneria tenuifolia (franseria) Tr.
Collomia gracilis (false gilia) Tr.
Heterotheca subaxillaris Tr.
Kallstroemia laetevirens Tr.
Lupinus sparsiflorus (lupine) Tr.
Martynia altheaefolia (small devil's-horns) 12
Mollugo verticillata (carpetweed) 324
Oenothera primiverus (evening primrose) 15
Opuntia discata (prickly pear) 15
Loeflingia pusilla Tr.
Lepidium lasiocarpum (peppergrass) Tr.
Plantago ignota (plantain) 818
Polygala puberula (milkwort) Tr.
Portulaca suffrutescens (purslane) Tr.
Prosopis velutina (mesquite) 1,570
Sida diffusa (spreading sida) 30
Solanum elaeagnifolium (742 fruits) (trompillo, prickly solanum) 156
Puffballs and fleshy fungi (undetermined) 12

Total species, exclusive of fungi, 41.It will be seen from Table 3 that while a large number of species of plants are represented in the totals from so many dens, a majority of them are of very minor importance, and that the seeds of grasses are the principal storage and probably therefore the principal food material. Six of the most important species of grasses (disregarding species furnishing less than 5 grams) comprise 85.6 per cent of the total weight of storage from 22 dens. Crowfoot grama (Bouteloua rothrockii) stands first in quantity in the total, forming 39.4 per cent of all stored material, 46 per cent of the six important grasses, and 45 per cent of all grasses. The largest amount of storage of any one species of grass in any one den on the Range Reserve also is of this species, 2,205 grams[5] (Table 1, den 1, p. 20, and Pl. VII, Fig. 2). This is exceeded by a dropseed grass, Sporobolus cryptandrus strictus, which amounted to 5,455 grams in a lot from Albuquerque, N. Mex. (Table 1, den 24, and Pl. VIII, Fig. 1).

Of the species other than grasses found stored in these dens, mesquite beans (Prosopis velutina) are most important both by weight and number of dens containing them. The total for the 22 Range Reserve dens is 1,570 grams, or 35.9 per cent of the seeds other than grasses, but only 5.1 per cent of the total storage. In bulk mesquite beans do not loom up large, as they are probably the heaviest material stored. Sections of pods which must have been dragged into the burrows are found, some of them certainly being much too long for carriage in the pouches. The species of plant other than grass found in the largest quantity in any one den, however, was Aplopappus gracilis, not recorded in quantity from any den until the excavation of the twenty-second, and then found in a very large bulk of soft, fluffy material, with most of the seeds separated from the heads, and weighing 1,030 grams (Table 1, den 22).

Any of the food materials above listed are likely to be found in the cheek pouches, while in addition such extraneous matter as stones and feces have also been found. All species of plants stored are accessible in the immediate vicinity of the mound, and when any particular plant is found seeding in abundance in the vicinity of the den it is likely to be represented in the storage. Usually the animals can be readily trapped with almost any kind of grain bait, as oats, rolled oats, rolled barley, and wheat; and nut meats also are attractive, though we have no record of the storing of any true nut in the dens, such not being available in the range of the animal on the Range Reserve.The following plants not represented in the list stored by the kangaroo rat on the Range Reserve have been found in the cheek pouches or mounds of spectabilis in other localities:

Amaranthus palmeri, Sesuvium portulacastrum, and Atriplex wrightii (alluvial soil of Santa Cruz Valley, Continental, Ariz., Bailey).

Cut leaves and stems of a small sagebrush (Franklin Mountains, Tex., Gaut).

Gutierrezia heads (San Juan Valley, N. Mex., Birdseye).

Verbesina enceliodes, Portulaca oleracea, Bouteloua gracilis, and Munroa squarrosa (Rio Alamosa, N. Mex., Goldman).

Tops of buds of Artemisia filifolia (Mesa Jumanes, N. Mex., Gaut).

Tumbleweed (Amaranthus graecizans), Russian thistle (Salsola pestifer), Munroa squarrosa, and Sporobolus cryptandrus strictus (Sandia Mountains, Albuquerque, N. Mex., Vorhies).


                                                                                                                                                                                                                                                                                                           

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